——— VOLUME 90, NUMBER 325 LIBRARY MAY 9, 1986 JUN 2 1986 HARVARD UNIVERSITY Neogene Paleontology in the northern Dominican Republic 3. The Family Poritidae (Anthozoa: Scleractinia) by Ann Budd Foster Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York, 14850 U.S.A. PALEONTOLOGICAL RESEARCH INSTITUTION Officers ei UREN ete ees LE P E M TT WILLIAM A. OLIVER, JR. A Че on boe Ce RR WILLIAM P. S. VENTRESS Ne LCS a CRIME NL LU DSS eme cto ME LM HENRY W. THEISEN рлык ыш ee ROBERT E. TERWILLEGAR PESSIS BARGE DRENSURERC o ei a ш кук ARE JOHN L. CISNE itane He ecc ТА RCM p T ET LU cu PETER R. HOOVER DROSUOGOUNSEE 225222222 00000022 - HENRY W. THEISEN Trustees BRUCE M. BELL (to 6/30/87) A. MCCUNE (to 6/30/86) E. ANNE BUTLER (to 6/30/88) CATHRYN NEWTON (to 6/30/88) RICHARD E. BYRD (to 6/30/86) WILLIAM A. OLIVER, JR. (to 6/30/86) JOHN L. CISNE (to 6/30/88) JAMES E. SORAUF (to 6/30/88) J. THOMAS DuTRO, JR. (to 6/30/87) ROBERT E. TERWILLEGAR (to 6/30/87) LEE B. 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Hoover Director Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York 14850 U.S.A. 607-273-6623 VOLUME 90, NUMBER 325 MAY 9, 1986 Neogene Paleontology in the northern Dominican Republic 3. The Family Poritidae (Anthozoa: Scleractinia) by Ann Budd Foster Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York, 14850 U.S.A. Library of Congress Card Number: 86-61076 Printed in the United States of America Allen Press, Inc. Lawrence, KS 66044 U.S.A. CONTENTS Page Abstract arten фатандар area ds tectam 47 Кезштпел sd. dpa ee РАН Г ҒА iat od 47 Introduction san: 48 Ackhowledementssesss Ike Niele ales клк dio Gener 49 Abbreviations of Repository Institutions ................. 49 Biostratigraphy and Paleoecology ........................ 50 Taxonomic Method Probo К е КОЛАТ of ТЕГ 56 МА ШАЙ та КЫ sooo ns uerbo domat тан SC TUR 59 ОҢДО н nennen las er лене davis quem 59 StatisticaleBroce dle wea. Pon de ныды ылы TUTO 63 Resultssandılnierpretationsumn 42020227. 68 General Comparisons with Other Faunas ............... 70 Systematic Paleontology Introduction ras sed ona spicae ee ORAN. sis WS Family Pontida: Gray 1842 а. 74 Сей ОРДе ИИ ШЕОЛ а о ar T3 Porites baracoaensis Vaughan, 1919 ............... 7/5) PORES EON VALORES а oo Мы 76 Porites macdonaldi Vaughan, 1919 ................ 78 Porites portoricensis (Vaughan, 1919) .............. 79 POLES Wayland BOME NON aere . 26s E . 81 Genus Goniopora Blainville, 1830 ................... Goniopora hilli Vaughan, 1919 22.2. er Goniopora imperatoris Vaughan, 1919 Goniopora calhounensis Weisbord, 1971 Genus Alveopora Blainville, 1830 .................... Alveopora tampae Weisbord, 1973 ................. Appendix Ia. Means of all corallite characters in the five species Ot: Pbritesdereinrdesoribedib а о oco a Appendix Ib. Means of all corallite characters in the three species of Gonioporasheremedesctibed iw a: SE ea Appendix Па. Canonical discriminant analysis of Porites groups o ed олан dtum Appendix IIb. Canonical discriminant analysis of Porites groups Кз ООУ eim etate ban alcala did tes Appendix IIc. Canonical discriminant analysis of Goniopora groups Банан aT Во anio as Appendix IId. Canonical discrimination analysis of five fossil and six modern Caribbean species of Porites .......... References Cited Index LIST OF ILLUSTRATIONS Text-figure Page 1. Map maicatme the location of the river sections sampled o errie.: e esis une ensure е 50 2. Bar charts summanzine the quantity of material collected ноа. 51 3. Diagrams showing the distributions of species within selected river sections .............с.................................... 52 4. Porites. Variation within species in corallite characters through a composite of the Rio Cana and Rio Gurabo stratigraphic EE IC ih 22.20 EU n ч e M E 54 5. Porites. Variation within species in colony shape through the composite stratigraphic section shown іп Text-figure 4 ............. DD 6. Goniopora. Variation within species in corallite characters and colony shape through the composite stratigraphic section shown in ТЕЧЕШЕ лн ЖЕСТ ктү ЗО PN сз oye eric UT нимо ни ременин peo шш 57 7. Scanning electron microscope photographs of modern Porites astreoides from different reef habitats near Discovery Bay, J amaica .... 58 8. Drawings showing боео charactersumeasuedew о нен н аии они н N 61 9. Serial acetate peels OWE Oe cotallitefofsBomtesfureatae. оинаи en IDA IO, ИЛР 62 10. Porites. Canonical discriminantanalysistot the NMB collections: онн en IAE ТӘ ARIAS 64 11. Goniopora. Canonical discriminant analysis of the NMB collections .......................................... eee ад 65 12. Porites subgroups. Canonical discriminant analysis of the NMB collections .................................................. 66 13. Goniopora subgroup. Canonical discriminant analysis of the NMB collections .............................................. ... 67 14. Means and standard deviations'for cight charactersam the five Poritesispecies .........................22222..2.22222 TRE 68 15. Means and standard deviations for eight characters in the three Goniopora species .................................2.м...... 69 16. Classification of Neogene Mediterranean types using the canonical discriminant analyses of the NMB collections ................ т 17. Porites. Canonical discriminant analysis of the NMB fossil collections and six modern Caribbean species ....................... 3/5) 18. Surface of a modern colony of Porites astreoides having a Synaraea-like appearance .......................................... 78 19. Surface of a modern colony of Porites astreoides with flaky septa ..........................................................ю 80 LIST OF TABLES Table Page 1. List of Neogene Caribbean poritid types used in statistical analyses ......................................................... 60 2. List of Neogene Mediterranean poritid types and modern Caribbean specimens used in statistical analyses ...................... 62 3. List and deseripbron of corallite characters analyzed ш Portes neun seen LO foldout inside back cover 4. List and description of corallite characters analyzed іп Goniopora .22......2.......2...................Х foldout inside back cover 5. Weighting of characters in the all Porites canonical discriminant analysis .................................................... 63 6. Weighting of characters in the all Goniopora canonical discriminant analysis ................................................ 67 7. List of all valid poritid species from the Miocene through lower Pliocene of the Caribbean, showing their current taxonomic Hub ан iss E POTN ed v ы Rt RI M RE d p E M m Am RE BS 74 NEOGENE PALEONTOLOGY IN THE NORTHERN DOMINICAN REPUBLIC 3. The Family Poritidae (Anthozoa: Scleractinia) By ANN BUDD FOSTER Geology Department The University of Iowa Iowa City, IA 52242 U.S.A. ABSTRACT Various multivariate statistical procedures are used to distinguish species in the reef-coral family Poritidae through a continuous Neogene sequence (five myr time interval) in the Cibao Valley of the northern Dominican Republic. Some older (by approximately 10 myr) material from the same region is also included in the analyses. The material consists of approximately 450 colonies (120 of which are measured) from 92 localities in four river sections. The colonies are first sorted into three genera, and approximately 30 characters measured on five calices per colony. The data are analyzed using cluster and canonical discriminant analyses to group the colonies into clusters representing species. Five species are so defined in Porites and three in Goniopora. These groupings are then used statistically to reclassify type specimens for 22 of the 25 described species of Neogene Caribbean poritids. Eight described species are thereby synonymized with four previously-described species in Porites and one new species of Porites, Porites convivatoris, n. sp., is discovered. Five described species are synonymized with two previously-described species in Goniopora. The stratigraphic range of three species of Porites and three species of Goniopora is also shown to extend back to the late Oligocene, thereby diminishing the significance of any presumed early Miocene adaptive radiation. Only one species was found to be endemic to the Dominican Republic and only one confined to the northern Caribbean. The rest are widely distributed throughout the Caribbean. Thus, the endemism previously believed common during the Neogene is shown to be far less extensive. Evolutionary trends within each species are preliminarily analyzed for various characters using nonparametric statistical procedures. In general, the results show that seven species experienced little or no evolutionary change (= stasis) through the sequence. Slight increases in corallite size are detected in two species, an increase in colony height in one species, and a more rounded colony shape in one species. These trends may be related to the general deepening of the environment; however, little correlation is found between lithology and morphology within species. Preliminary analyses of the relationship between intra- specific variation and poritid abundance and diversity yield significant results, suggesting that intraspecific trends may be environmental and that future study of coral species associations may offer insight into paleoenvironmental interpretations. Statistical comparisons with the Miocene Mediterranean poritids show that no species co-occur in the two provinces during the Neogene. Similarly, none of the studied Neogene species of Porites resemble modern Caribbean species of Porites, signifying that all nine poritid species studied must have become extinct and the modern Caribbean species of Porites radiated during the late Pliocene or early Pleistocene. This study represents part of a multidisciplinary project on the stratigraphy of the northern Dominican Republic, coordinated by P. Jung and J. B. Saunders of the Naturhistorisches Museum Basel, Switzerland. RESUMEN Se usan varios procedimientos estadisticos multivariados para distinguir especies de la familia de corales arrecifales Poritidae a travéz de una secuencia Neogéna continua (intervalo de tiempo de cinco millones de anos) en el valle Cibao, al norte de la Repüblica Dominicana. Algun materiel de mayor edad (aproximadamente 10 millones de afios) de la misma region es tambien incluyendo en las analises. El material consiste en aproximadamente 450 colonias (120 de las cuales son medidas) de 92 localidades en cuatro secciones del rio. Primero las colonias son divididas en tres géneros y aproximadamente unos 30 caracteres son medidos en cinco cälices por colonia. Los datos se analizan usando anälisis de grupos у anälisis canónicos discriminativos para agrupar las colonias en grupos representantes de especies. Cinco especies se definen asi en Porites y tres en Goniopora. Estas agrupaciones son luego usadas estadísticamente para reclasificar clases de especímenes de 22 de las 25 especies descritas de poritidos Neogénos del Caribe. Ocho especies descritas son comparadas análogamente con cuatro especies previamente descritas en Porites, y una nueva especie de Porites, Porites convivatoris, n. sp., es descubierta. Cinco especies descritas son comparadas análogamente con dos especies previamente descritas en Goniopora. Se ha demonstrando que el alcance estratigráfico de tres especies de Porites y tres especies de Goniopora se extiende hasta el Oligoceno tardío, disminuendo de ese modo, la importancia de la presumible radiación temprana adaptiva Miocena. Se encontró que solo una especie es endemica de la República Dominicana y que solo una especie fue limitada al norte del Caribe. El resto está distribuido ampliamente a travéz del Caribe. Por lo tanto, la endemia que previamente se consideró común durante el Neogéno, ha demonstrando ser mucho menos extensa. Las tendencias evolutivas dentro de cada especie son analizadas preliminarmente por varios caracteres usando procedimientos estadísticos no paramétricos. En general, los resultados muestran que siete especies experienciaron poco a ningún cambio evolutivo (estásis) a travéz de la secuencia. Se detectan pequeños aumentos del tamaño de los esqueletos de pólipos en dos especies, un aumento de la altura de la colonia en una especie, y una forma mas redondeada de la colonia en una especie. Estas tendencias podrían estar relacionadas con la profundización general del medio ambiente, sin embargo, se encontró poca correlación en una 48 BULLETIN 325 escala mas pequeña entre litologia y morfología entre las especies. Análisis preliminares de la relación entre variación intraespecifica y abundancia y diversidad porítida producen resultados importantes que sugieren que las tendencias entraespecíficas podrían ser ambientales y que el estudio futuro de las asociaciones de las especies de corales podría ofrecer un entendimiento de las interpretaciones paleoambientales. Comparaciones estadísticas con los poritidos Mediterráneos del Mioceno muestran que ninguna especie co-ocurre en las dos provincias durante el Neogéno. En forma similar, ninguna de las especies Neogénas estudiadas de Porites se parece a las especies de Porites del Caribe moderno, lo cual significa que todas las nueve especies referidas ya estudiadas deben haberse vuelto extinctas y que las especies de Porites del Caribe moderno fueron radiadas durante el Plioceno tardío o Pleistoceno temprano. Este estudio representa parte de un proyecto multidisciplinario de la estratigrafía del norte de la República Dominicana coordinado por P. Jung and J. B. Saunders, Naturhistorisches Museum Basel, Switzerland. INTRODUCTION General surveys of taxonomic distributions suggest that a major extinction event occurred in the evolution of Caribbean reef-corals during the late Oligocene. This event was followed by a smaller radiation of taxa in the early Miocene. Many modern coral taxa are be- lieved to have originated during this radiation, together with numerous endemic short-lived Neogene corals that gradually became extinct during Pliocene and ear- ly Pleistocene time (Frost, 1977a, 1977b). However, the detailed evolutionary patterns of the radiation and subsequent decline are poorly documented, largely be- cause the definition and evolutionary patterns of many species are based only on a few, poorly-preserved spec- imens. In addition, the stratigraphic relationships be- tween many coral faunas are known only on a gross scale involving the correlation of several widely-sep- arated sections. The purpose of the present investi- gation is to redefine Neogene coral species using ma- terial collected at regular stratigraphic intervals through a long, continuous sequence. The species redefinitions are made by analyzing morphologic variation in groups of specimens. Emphasis is placed on tracing evolu- tionary patterns within each species in detail and using the results of this analysis to postulate possible evo- lutionary relationships. The present study is prelimi- nary in nature and is intended to provide the basis for a more complete systematic revision of all Caribbean Tertiary corals. This paper is the first in a series of coral faunal descriptions of the middle Miocene to middle Pliocene of the northern Dominican Republic. The coral col- lections were made between 1978 and 1980 by J. Geis- ter, P. Jung, J. B. Saunders and coworkers as part of their large-scale multidisciplinary project on the pa- leontology and stratigraphy of the Neogene of the Ci- bao Valley region, and all collecting localities are keyed into their detailed stratigraphic sections (Saunders er al., 1982; Saunders, Jung, and Biju-Duval, 1986). The project involves specialists on a wide variety of taxo- nomic groups working on material from the same lo- calities. Therefore, coral evolutionary patterns can eventually be compared with those of other taxa, and ecological and stratigraphic relationships can be inter- preted using a large comprehensive data set. Micro- fossil dates are also available for almost all localities. The collections themselves were made in bulk at reg- ular stratigraphic intervals, offering a rare opportunity to trace the evolution of large coral populations through a five-million year sequence. The northern Dominican Republic was selected for study because it represents one of the longest, most continuous, and best-studied sections through Neo- gene coral deposits in the Caribbean. The corals are also exquisitely preserved. This first coral paper in the series covers the most abundant coral family, the Pori- tidae. Two genera in this family are discussed in detail, one of which, Porites Link, 1807, represents an essen- tial component of modern Caribbean and Indo-Pacific reefs. The other, Goniopora Blainville, 1830, became extinct in the Caribbean during the late Pliocene and is represented today only by Indo-Pacific forms. A third poorly-represented genus, A/veopora Blainville, 1830, which also occurs today only in the Indo-Pacific and is rarely reported in Caribbean Tertiary deposits, is briefly discussed. Previous work on the systematics of the Dominican Republic corals has involved only a portion of the fauna and was based on limited material. The first major study describing the Neogene corals (Duncan, 1863, 1864, 1868) was based on the Heneken collec- tion (Heneken, 1853), which is now deposited at the BM(NH). In these publications, Duncan described 48 species (32 of which were new) from the “Nivajé Shale” and the “Silt of the Sandstone plain”. Most of these descriptions were based on single specimens or frag- ments of specimens and were re-interpreted by Vaughan (1919) to represent a total of 28 species, 20 of which were new. Vaughan found only two species described by Duncan to belong to the family Poritidae, Alveopora fenestrata Dana, 1846 and Porites collegniana Mi- chelin, 1842. Shortly after Duncan’s publications, Pourtalés (1875) compiled a list of corals collected by W. B. Gabb (Gabb, 1873; deposited at MCZ and ANSP). The list included 30 taxa, five of which could not be assigned to any species. Only one species be- longed to the family Poritidae, Porites furcata La- marck, 1816. DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 49 Almost fifty years after these initial works, Vaughan (1919, p. 217) listed 28 coral species in the Maury collection (Maury, 1917; deposited at USNM) from the Neogene of the Dominican Republic. One of these species, Agaricia dominicensis Vaughan, 1919, he for- mally described as new, and six others were cited as new and to be described in a later publication, which never appeared. Two of these six species apparently belonged to the Poritidae, and their descriptions are given in an unpublished manuscript on the Maury col- lection by Vaughan (date unknown). In 1919, Vaughan and Cooke made large, well-documented collections of well-preserved coral specimens in the Dominican Republic, mainly from Rio Yaque del Norte, Rio Mao, Rio Gurabo, and Rio Cana. These are listed by USGS locality number in Vaughan et al. (1921, pp. 105-168) and deposited at the USNM. Formal systematic de- scriptions of this material were never completed. Nine new species were eventually described in Vaughan and Hoffmeister (1925); however, these were based pri- marily on older material from the Gabb collection and included no species belonging to the Poritidae. Since Vaughan's work, very little has been done on the Neogene coral fauna of the Dominican Republic. Ramírez (1956) has listed five Neogene coral species from Rio Amina of the northern Dominican Republic, and Ricart y Menéndez (1981a, 1981b, 1983) has de- scribed skeletal structures in three species of Antillo- cyathus Wells, 1937, from an undisclosed locality in the Dominican Republic. Neither worker has attempt- ed to describe the poritids. ACKNOWLEDGMENTS This work was made possible by a six-month inter- national postdoctoral fellowship from the Swiss Na- tional Science Foundation (No. 88.143.0.83) for re- search at the Naturhistorisches Museum Basel (NMB). I would especially like to thank Dr. Peter Jung for his advice and guidance during the tenure of the fellowship and Mr. John B. Saunders for his help with all aspects of Caribbean stratigraphy. I am very grateful to a num- ber of people in Basel for their technical assistance: W. Suter for making the whole-colony and surface pho- tographs; К. Müller for preparing thin-sections; С. Scherler for library assistance; С. Lüönd and К. Gug- genheim for scanning electron microscopy; D. Ber- noulli and М. Düggelin for help with thin-section pho- tography; H. Moser, H. Christen, and P. Schmid for Computer advice; and R. Panchaud for assistance with Curation of specimens. I thank the following individuals and institutions for arranging loans of museum material during the tenure Of the fellowship: S. D. Cairns and К. Rützler of the U. S. National Museum of Natural History (Washing- ton, DC), M. B. Best of the Rijksmuseum van Na- tuurlijke Historie (Leiden, The Netherlands), G. E. de Groot of the Rijksmuseum van Geologie en Mine- ralogie (Leiden, The Netherlands), B. R. Rosen and P. Cornelius of the British Museum (Natural History) (London, United Kingdom), M. Grasshoff of the Na- tur-Museum Senckenberg (Frankfurt, West Germany), C. C. Jones of the Academy of Natural Sciences of Philadelphia (Philadelphia, PA), R. L. Langenheim, Jr. of the University of Illinois (Urbana, IL), S. Stuenes of Uppsala Universitet (Uppsala, Sweden), R. C. Eng of the Museum of Comparative Zoology (Cambridge, MA), N. Eldredge of the American Museum of Natural History (New York, NY), W. D. Hartman of the Pea- body Museum of Natural History (New Haven, CT), O. Elter of the Museo di Zoologia Sistematica (Torino, Italy), O. Schultz of the Naturhistorisches Museum Wien (Vienna, Austria), and S. Wise of Florida State University (Tallahassee, FL). I also thank J. Maréchal for assistance in locating the Lamarck specimens at the Muséum d’Histoire naturelle in Paris (France); M. D. Serrette for photographing the Paris material; and J. Maréchal and S. Barta-Calmus for assistance with the Chevalier collections at the Institut de Paléontologie in Paris. I am grateful to J. Geister (Bern, Switzerland) for introducing me to the NMB coral collections and for help with initial sorting of the corals and X-radiog- raphy; to B. R. Rosen (London, United Kingdom) and J. W. Wells (Ithaca, NY) for their lengthy discussions of poritid morphology and taxonomy; to G. Klapper (Iowa City, IA) for advice on zoological nomenclature; and to A. H. Cheetham (Washington, DC) for discus- sion of statistical techniques. The following provided technical assistance: T. Bahns in preparation of thin- sections and acetate peels; U. Dogan in scanning elec- tron microscopy; T. Druecker for translations; L. John- son and R. Petrick in typing; J. Klapper in preparation of plates. I thank S. D. Cairns and J. W. Wells for reviewing the manuscript. J. Geister, J. Golden, P. Jung, and J. B. Saunders also generously read the manuscript and offered many helpful suggestions. Additional funds were provided by a grant from the U. S. National Science Foundation (BRS83-07109). ABBREVIATIONS OF REPOSITORY INSTITUTIONS AMNH: American Museum of Natural History, New York, NY, U.S.A. ANSP: Academy of Natural Sciences of Philadelphia, Philadelphia, PA, U.S.A. BM(NH): British Museum (Natural History), London, England, United Kingdom FSU: Florida State University, Department of Geol- ogy, Tallahassee, FL, U.S.A. MCZ: Museum of Comparative Zoology, Harvard University, Cambridge, MA, U.S.A. MHNP: Muséum National d’Histoire naturelle, Paris, France NHMW: Naturhistorisches Museum Wien, Vienna, Austria NMB: Naturhistorisches Museum Basel, Switzerland PIU: Paleontologiska Institutionen Uppsala, Uppsala, Sweden SUI: State University of Iowa, Iowa City, IA, U.S.A. UCMP: University of California Museum of Paleon- tology, Berkeley, CA, U.S.A. UI: University of Illinois, Department of Geology, Ur- bana, IL, U.S.A. USGS: United States Geological Survey, Washington, DC, U.S.A. USNM: U. S. National Museum of Natural History, Washington, DC, U.S.A. (five-digit catalogue num- bers refer to the Department of Invertebrate Zool- ogy, six-digit catalogue numbers refer to the De- partment of Paleobiology) YPM: Yale Peabody Museum, New Haven, CT, U.S.A. BIOSTRATIGRAPHY AND PALEOECOLOGY The Poritidae are abundant and widespread in four of the collected river sections (Rio Cana, Rio Gurabo, Rio Mao, and Rio Yaque del Norte) of Saunders, Jung, and Biju-Duval (1986) through the Neogene of the BULLETIN 325 Cibao Valley (Text-fig. 1). They were not found else- where in the study area. Specimens were collected at a total of 92 localities ranging in age from late Oligo- cene to middle Pliocene. Two species, Porites porto- ricensis (Vaughan, 1919) and P. baracoaensis Vaughan, 1919, are especially prolific, occurring in more than 30 localities each (Text-fig. 2a). Porites portoricensis was found in all four river sections (Text-fig. 3), whereas P. baracoaensis occurs in all but the older, more con- glomeratic Rio Yaque del Norte section. The three remaining species of Porites (P. waylandi, nom. nov., P. convivatoris, n. sp., and P. macdonaldi Vaughan, 1919) were collected in 15 or more localities each (Text-fig. 2a). Of these three species, P. waylandi was found in all but the Rio Mao section. It appears to be more common in the Rio Yaque del Norte section and the lower parts of the Rio Gurabo and Rio Mao sections. Porites convivatoris is restricted to the Upper Miocene and Lower Pliocene of the Rio Cana and Rio Mao sections. Porites macdonaldi occurs most com- monly throughout the middle portions of the Rio Gu- rabo section and has only a limited distribution along Rio Cana and Rio Mao (Text-fig. 3). Goniopora Blainville, 1830 and Alveopora Blain- ville, 1830, were less commonly found (Text-fig. 2a). Of the three species of Goniopora, G. hilli Vaughan, 1919 occurs most frequently. It was found in all but the Rio Mao section and appears to be restricted to Upper Miocene (possibly lowermost Pliocene) and old- er sediments. Goniopora imperatoris Vaughan, 1919 N - 10 20km 1 Rio Cana 2 Rio Gurabo GUAYUBIN CT B | L_J Upper Cenozoic E N A О (5:1 Oligocene - Early Miocene ? 5 Canada Zalaya » 6 Rio Yaque del Norte 3 4 A Ме 7 City of Santiago Ф 4% 4 4 8 Arroyo Puñal © N 8 & > 9 Rio Verde 9 x 8 ы 3 - 1 2 5 VALVERDE ESPERANZA NAVARRETE EN. ZAMBA $ МАО ? 5 BEN [UNS 7038 Кеш, En LOSQUEMADOS & Uy, SANTIAGO 8 22 RODRIGUEZICT— 4 ny ES ann У, [SANTIAGO MOCA 6 8 C BAITOA Text-figure 1.—Map indicating the location of the river sections sampled. Members of the Poritidae were found only in four sections: (1) Río Cana, (2) Río Gurabo, (3) Río Mao, and (6) Río Yaque del Norte (map from Saunders, Jung, and Biju-Duval, 1986). DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 51 A 30- Y E c a ә Ei ті 9 c < a Е 204 О N е 1 о - + i 2 c z = т - т 2 = ч Ы 104 © 5 ; 5 2 o z ш е в T QU W a exo S «ж ой? © өм o? „се ве \0 yor a oN A o oo" oc? N RS) có \ «e*t 2504 4 = 200 - = | 3 | O | O Qz Lu сп = 100- > z j x 5 50- = | | | | j 9 229 yor go? AÑ х n e e o BER See cot e e one“ Text-figure 2.— Bar charts summarizing the quantity of material collected. (A) Percentage of all Poritidae localities containing each species. “п” = total number of localities represented. (В) Total estimated number of colonies collected of each species. 52 BULLETIN 325 occurs in all but the Rio Gurabo section and seems to RIO GURABO be restricted to sediments of the same age as G. hilli. Goniopora calhounensis Weisbord, 1971 was found only at two localities (one on Rio Cana and the other on Rio Gurabo), both of which are relatively high in the section (Text-fig. 3). Alveopora only rarely occurs in the region studied and is represented by one species (A. tampae Weisbord, 1973). It is confined to the Up- per Oligocene Tabera Group (Text-fig. 3b). The overall abundance of the seven common poritid species in the NMB collections cannot be determined with any accuracy. Numbers of individuals (i.e., col- onies) cannot be directly counted with certainty, be- cause the specimens are so severely fragmented. Vol- umetric estimates do not reflect the genetic structure of the population, because of extensive differences in 0 3s adult size and reproduction (especially, the relative gore grace contribution of asexual vs. sexual reproduction) be- tween coral species. These difficulties are best dem- RIO CANA onstrated by the three following examples. First, spec- Bo te imens ofthe two branched species (P. portoricensis and o P. baracoaensis) consist of numerous broken frag- E 3 ments, from which it is impossible to reconstruct the a 1000] x% Ж original colonies. In such cases, it is conceivable that all the pieces of a species from one locality belonged to the same colony or at least the same clone. Second, even large, massive corals such as G. hilli are often so badly broken that it is difficult to determine which pieces belonged to the same colony. Third, P. convi- vatoris, which formed small, compact colonies, is usu- ally preserved unbroken and is represented by numer- ous countable individuals at several localities (e.g., 83 colonies at NMB loc. 16928). Because of its small size, however, P. convivatoris is not volumetrically impor- tant in any NMB sample. In light of these problems, to describe abundance, G the number of colonies of each species has only been subjectively estimated, and the general results are RIO YAQUE DEL NORTE graphed on Text-figure 2b. Porites convivatoris is rep- AM resented by the greatest number of colonies and the two species of Goniopora by the least. The remaining four species of Porites have roughly equivalent values, with P. waylandi possibly having the fewest individ- 8 а 3 1000 4 > m о 8004 <1 Mao 6004 | Early Pliocene | Middle Pliocene 400 - 5 š thickness in meters Gurabo 2004 | Late Miocene S го Сегсадо asis üf anê do ад” il ensis N аточ" o" м mas! a > —0 1200 4 n-8 п-9 1 4 - x2 8004 n=l Mao 600 + | Early Pliocene thickness in meters Gurabo 4004 ў »— ж ж py D м< ne - en гә - ee 200 - Cercado Late Miocene 2 ف اي‎ з \Ч N 9 کاو‎ Ci Сы ONT eot eet voco? об? 09 со“ کې‎ on ° с 00 Late Miocene т Е | Q SIT A > 1 xS xa Pac thickness in meters (oped uals. Study of patterns of morphologic variation across a composite of the Rio Cana and Rio Gurabo strati- graphic sections (spanning a time interval of approx- imately five myr) shows pronounced stability within each species (Text-figs. 4, 5, 6). Variability within a o L a a - Baitoa Early - Middle Miocene Text-figure 3.—Diagrams showing the distributions of species within selected river sections. “п” = total number of localities con- o4 taining each species. Numbers to the right of points along each vertical distribution line indicate the number of localities represented by each point. (A) Río Gurabo, (B) Río Cana, (C) Río Yaque del Norte. LOPEZ -- 1 Tobera censis » oyien d" [me oratoris e [Oligocene porto DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 53 species is indeed high within many stratigraphic in- tervals, but overall change upsection within each species is only slight or nonexistent. This general result is con- firmed by a Runs statistical test and a Binomial sta- tistical test performed on 50-m and 100-m interval time series data for each species using the SPSS-X NPAR procedure (SPSS Inc., 1983). In all five species of Porites, canonical variable 1 (a variable describing corallite wall structure and corallite spacing; see de- scription of all Porites canonical discriminant analysis in the next section) remains unchanged upsection (Text- fig. 4a). Results of regression analysis and Spearman’s rank order correlation coefficients (SAS Inst., 1982, PROC REG and PROC CORR) indicate that no re- lationship or correlation exists between relative strati- graphic position and this canonical variable. However, canonical variable 2 of the same canonical analysis (a variable describing the inverse of corallite size) does appear to decrease upsection in P. waylandi (rg = —0.750, p = 0.012). Values of canonical variable 2 for the remaining four species (Porites portoricensis, P. baracoaensis, P. convivatoris and P. macdonaldi) all zigzag back and forth upsection without any detectable Overall trend (Text-fig. 4b). Colony shape also does not appear to change upsection. Spearman’s rank order correlation coefficients show insignificant correlations between stratigraphic position and branch thickness in the two branching species of Porites (Text-fig. 5a), and between stratigraphic position and the ratio of colony height to length in the three massive species of Porites (Text-fig. 5b). In the two species of Goniopora, signif- icant directional changes occur upsection only in col- Ony shape (rs = 0.580, p = 0.002) of G. imperatoris (Text-fig. 6) and in colony height of G. hilli (ra = —0.717, P = 0.0004). Significant differences in canonical vari- able 1 (a variable describing corallite size; see Gonio- bora canonical discriminant function analysis in the next section) occur also between specimens of G. hilli Collected on Rio Yaque del Norte and those collected оп Rio Cana and Rio Gurabo using two nonparametric rank sum statistical tests, the Wilcoxon 2-sample test (р = 0.0414) and the Kruskal-Wallis test (р = 0.0311), of the SAS NPARIWAY procedure (SAS Inst., 1982). This general stability suggests not only a lack of evo- lutionary development but also a lack of response to environmental change, since the Cibao Valley Neogene Section is believed to have been deposited under grad- ually deepening conditions (Saunders, Jung, and Biju- Duval, 1986). It is unclear if the slightly increased Corallite sizes in P. waylandi and G. hilli, and larger, more rounded colony shape in G. imperatoris and G. hilli are caused genetically or environmentally. Spear- man’s rank order correlation coefficients show that li- thology is not simply correlated with canonical vari- able 1, canonical variable 2, or colony size or shape in any of the examples above. This lack of correlation may be partially related to the fact that many of the corals in this study were found in transported rubble layers within silts. However, significant correlations with the diversity and abundance of poritid species collected at each locality do exist in some instances, suggesting that some intraspecific trends may be en- vironmental. For example, canonical variable 2 (a variable describing corallite size; see description of all Porites canonical discriminant analysis in the next sec- tion) is positively correlated with abundance (i.e., number of poritid colonies collected per locality) in P. waylandi (rs = 0.700, р = 0.004). Decreases in thick- ness of plates of P. macdonaldi (rs = —0.532, р = 0.041) and more rounded shapes in P. waylandi (rg = 0.512, p = 0.051) appear correlated with increased number of species. Flatter, more encrusting colonies are correlated with increased diversity (calculated us- ing the Simpson diversity index as given in Pielou, 1969) in both G. hilli (rg = —0.790, p = 0.0001) and С. imperatoris (rs = —0.565, p = 0.0001). Assuming that abundance and diversity of the collected poritids are related to reef development, these preliminary re- sults suggest that study of variation in some massive species (especially in colony size and shape) may be potentially useful in interpreting subtle changes in en- vironment upsection. The details of this relationship, however, can only be further explored when occurrence data are compiled for more taxa through the studied stratigraphic sections. Distribution patterns for occurrences of single species offer less paleoecological potential. The collected po- ritid species do not appear to be confined to any one lithology, although some limitations have been ob- served. Porites baracoaensis does not occur in the con- glomerates at the base of the series, whereas P. way- landi and the two abundant species of Goniopora occur more commonly there. The restriction of P. conviva- toris to a few isolated pockets in the silts of Rio Cana and Río Mao is especially enigmatic and may be related to reproduction, recruitment, its possible symbiont, or other biological factors unrelated to the physical en- vironment. Few paleoecological interpretations can be deduced by directly comparing these fossil species with modern analogs. None of the poritid species collected during the study exist today or have direct modern counter- parts (see p. 71). All three genera occur today in a wide variety of reef habitats. Porites, especially, is widely distributed across modern reefs and occurs from shal- low nearshore zones across the lagoon and reef crest into the deep forereef areas. Furthermore, with the exception of P. convivatoris, С. calhounensis, and Al- 54 BULLETIN 325 їй m 12004 Sue aa 3 а 7007 ] : Р RC 4 10004 i ич ар, же " 8007 E A ] Sepp —1— Eb R = R ] : l П I ] А о 0 1 = 500 4 6007 y б 1 3 he A 1 а 3pm 1 5 R t N 2 qm E A ^ 1 Mg =a B 4007 : z о J 4 -+—5— 3 4004 393 -—————— ] A 300- ны ng T pe 1 ыл 20074 ] 5 100- | ОР куык e аа бы = T $ Eh y T M А m me -8 -4 0 4 8 А CANONICAL VARIABLE 1 m m А 3 700 1 : 1 en 2 - 10004 == i 2 600 7 : e^ : m B00-] INN - cc oc Б ар" =e эсе ш R сі A тИ Ул [ I ] er, [9] 0 1 N... 500 - 80054 ағалы ea nite ceolod ЛШ ТА 5 Те 1 RA UU 2 3 ЖТ e | Ем А А ] А” B AGUN " 412 " о dl ] سإ‎ 4. E b 400: 393 Zum quu “йр ae Glee? کد‎ міні 1 +; I N 3004 аы 1 24 Чч a 1 dh 2007 , 4 PT 1 MA + 04 : Т е рт в -4 -2 0 г 4 CANONICAL VARIABLE 2 Text-figure 4.— Porites. Variation within species in corallite characters through a composite of the Rio Cana and Rio Gurabo stratigraphic sections. The composite was constructed using the Rio Cana section (Saunders, Jung, and Biju-Duval, 1986, text-fig. 16) as a standard and scaling the Rio Gurabo section (Saunders, Jung, and Biju-Duval, 1986, text-fig. 6) relative to it. To accomplish this, three exact elevations were correlated between the two sections: (1) the unconformity with the Tabera Group at the base of each section, (2) the boundary between the Globorotalia humerosa and G. margaritae foraminiferal zones, and (3) the boundary between the G. margaritae and G. miocenica zones. The positions of localities within each section were determined using text-figures 4 and 15 of Saunders, Jung, and Biju-Duval (1986). The points (labelled 1-5) on the diagram above represent means for every 100-meter interval along the composite section. 1 = Р. portoricensis, 2 = P. baracoaensis, 3 = P. waylandi, 4 = P. convivatoris, 5 = P. macdonaldi. Horizontal lines on either side of each point are one-half standard deviation in length. (A) Canonical variable 1 of the all Porites canonical discriminant analysis. As described in the following section, DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 55 m m 1200 Arie) = Rm RE 7004 2 4 1000 en adus 6007 ai! 800 m De 2 ү R E R 2: | I S : 0 0 2 Б 500 - 600 i г) (8 ; Т А 2 | R 1 N = A A ES, B 400 0 el 4004 393 E e e max.s.d. 3004 Se 200 ad 2 2007 100- | (A eee 0 T T T d ы Т T 0 20 40 60 A BRANCH THICKNESS (MM) an 12004 stk == Ey E ] 5 roo] ] ] 5 4 1000-4 5 | 6007 4 | воо- de 1 | 5 R “ R 1 5 | I 1 È | (0) 0 1 5; 500 + 6004 d G 6 ] Ë U A 1 ? 3 2; Б | А Je вәмітпеіеы «65 Yaris A A dud) AR тыш DEPO AA — UI B 400-4 max. s.d. о ] 5 e 002.393 TET : us 8 300 1 dal | 200-4 es 4 2004 ] f 1004 ] 1 S etu s de сы 9 f e T r сне T VEL T > eE B 0.0 055 1*0 5 COLONY HEIGHT / COLONY LENGTH Text-figure 5.— Porites. Variation within species in colony shape through the composite stratigraphic section shown in Text-figure 4. 1 = Ti portoricensis, 2 = P. baracoaensis, 3 = P. waylandi, 4 = P. convivatoris, 5 = P. macdonaldi. The lengths of the maximum standard deviation are given. (A) The three massive species of Porites. (B) The two branching species of Porites. ———— Wall structure and calice elevation are most heavily weighted on this canonical variable. (B) Canonical variable 2 of the all Porites canonical discriminant analysis. The inverse of corallite diameter is heavily weighted on this canonical variable. The lengths of the maximum standard deviation are given. 56 BULLETIN 325 veopora tampae, all the poritids in the study have wide biogeographic distributions across the Caribbean throughout Neogene time. However, the local distri- bution patterns and morphologic variation have not been studied adequately at other Neogene Caribbean localities (especially the type localities for these species) for use in interpreting the patterns observed in this study. In conclusion, because of the wide ecologic and bio- geographic distributions, extensive variability, and the apparent stability of most of the poritid species in the collections of Saunders, Jung, and Biju-Duval (1986), it seems premature to assign to them any biostrati- graphic or paleoecologic importance. Their long du- rations may eventually prove detrimental to their potential biostratigraphic value, and their wide envi- ronmental distributions may decrease the paleoeco- logical value of studying the occurrences of individual species. However, preliminary results suggest that (1) patterns of variation within single species and (2) pat- terns of abundance and diversity for the complete coral fauna may eventually provide important keys to pa- leoenvironmental interpretation of the sections stud- ied. Since none of fossil species studied have direct modern counterparts and the variability within species does not appear clearly to be correlated with lithology, the true value of studies of intraspecific variation can- not be determined until all the corals in the collections of Saunders, Jung, and Biju-Duval (1986) have been studied taxonomically, and data are available on the distribution and abundance of the complete coral fau- na. Interpretations of overall patterns of diversity and abundance for the coral fauna would require: (1) com- parisons with other Neogene and modern Caribbean reefs, and (2) substantiation of patterns by analyzing the spacing, size, and arrangement of in situ coral col- onies at selected localities, as well as by determining whether colonies at these localities are preserved in growth position. Since modern corals appear distrib- uted across reefs in zones composed of unique asso- ciations of species (see Geister, 1983), one further pos- sible method for future environmental interpretation using data for the entire coral fauna in the collections of Saunders, Jung, and Biju-Duval (1986) would in- volve quantitative analyses of associations of species [for example, by procedures similar to reciprocal av- eraging and correspondence analysis as used by Cisne and Rabe (1978)]. TAXONOMIC METHOD PROBLEM The Poritidae include many of the most abundant reef-corals throughout the world, since early Neogene time. Yet, because of their extensive variability and relative lack of distinct skeletal features, their taxon- omy is highly problematic. As explained by Bernard (1901, 1903, 1905) and reconfirmed by Brakel (1976), the species do not form discrete morphologic clusters but show a gradational continuum in all morphologic characters. This apparent lack of distinct groupings led Bernard to abandon the species concept and Linnean taxonomy altogether (see detailed discussion in Cock, 1977), and it led Brakel to identify only “phenons” in his morphometric study of corallite variation in Ja- maican representatives of Porites Link, 1807. In modern Caribbean Porites, various interpreta- tions have been proposed to explain the number of species that exist at any one locality (Pl. 15). Several workers have recognized only two species, one massive (P. astreoides Lamarck, 1816) and the other branched [P. porites (Pallas, 1766)] (see Vaughan, 1901; Cairns, 1982; Roos, 1971; Almy and Carriön-Torres, 1963; Zlatarski and Estalella, 1982). In doing so, they typi- cally synonymize the three branching forms P. porites, P. furcata Lamarck, 1816, and P. divaricata Lesueur, 1821. Doubt is also expressed as to the validity of Porites branneri Rathbun, 1887, which is sometimes believed synonymous with P. astreoides Lamarck, 1816 by workers in the Caribbean (e.g., Roos, 1971) but appears to represent a distinct morphologic group in Brazil (Laborel, 1969). Although some general relationships between colony shape and environment have been observed (Roos, 1967), variation in calice morphology within species is widespread and does not follow any consistent pat- tern that can be correlated with environmental vari- ables. In Porites furcata, for example, skeletal struc- tures appear more heavily calcified, calices shallower, and wall reticula thicker with increased depth on the forereef near Discovery Bay, Jamaica. However, in Porites astreoides, calice elevation is deepest and skel- etal structures thinnest in low-light environments. Cor- allite diameter is largest in the backreef and smallest on a nearshore patch reef, both representing shallow, high-light environments with equivalent wave energies (Text-fig. 7). This lack of correlation between calice Text-figure 6.—Goniopora. Variation within species in corallite characters and colony shape through the composite stratigraphic section shown in Text-figure 4. 1 = G. hilli, 2 = G. imperatoris. (A) Canonical variable 1 of the all Goniopora canonical discriminant analysis. As described in the following section, corallite diameter is most heavily weighted on this canonical variable. Horizontal lines on either side of each point are one-half standard deviation in length. (B) Canonical variable 2 of the all Goniopora canonical discriminant analysis. Septal length and elevation are most heavily weighted on this canonical variable. Horizontal lines on either side of each point are one-half standard deviation in length. (C) Colony shape. The lengths of the maximum standard deviation are given. DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 37 m ee ee 7007 4 1000-4 600 8004 Қ 1 2 al | I Ser 0 0 500 4 6004 б D U A R 3j N met кашу A A B 4004 o سا‎ OO 3007 Р 2007 E 100- e A of T T T T A -4 -2 0 2 4 CANONICAL VARIABLE 1 m 1200-] pli) ecce тоо] ] d 1000-4 6007 800-4 R 4 R ] i I 3 о 0 5004 5004 6 (9 U A R x N 3 A A 1 B 400-] о 1 zi 2 ЧОО зун mm 3004 | 2007 2007 1007 | 0--------- 0 т т T T T | B ES -2 -1 0 4 а CANONICAL VARIABLE 2 max. s.d > 12004 ] a ж ere | 700- 1 2 4 10004 6004 Р 800-4 B R 5] R | 1 о 0 5004 5007] 6 0 U A R N A | A ] B 400-4 0 2 | 4004 393 ee 3007 ; 2004 y 200 1 100 ee 9 T T ы i 3 с 0.0 0.2 0.4 0.6 0.8 1.0 COLONY HEIGHT / COLONY LENGTH 58 BULLETIN 325 morphology and environmental variables contrasts with the strong correlations found in species of Mon- tastraea Blainville, 1830, and Siderastrea Blainville, 1830, collected at the same localities (Foster, 1979, 1980). To deal with such complex variation in Porites, Vaughan (1907) described 16 forms of Hawaiian Po- rites compressa Dana, 1846 and presented a complex diagram to illustrate their interrelationships. He did not, however, attempt to explain the cause for this variation. Work on reproduction in modern Porites suggests that much intraspecific variability may be directly caused by high genetic variation. In general, Porites is especially fecund and reproduces sexually by planu- lation (Goreau, Goreau, and Hayes, 1981; Kojis and Quinn, 1982). Therefore, a species would not neces- sarily form a completely coherent morphologic cluster but could include numerous scattered outliers, whose morphologies cannot be explained without knowledge of the underlying genetic controls. This interpretation of widespread genetic variation is also supported by Brakel (1977) in his morphometric analyses of modern Porites skeletons. Given these difficulties in delineating modern species, a relatively objective, statistical approach has been adopted in the present analysis of fossil species. Spec- imens were subjectively sorted into genera and grouped into clusters using various multivariate analyses of morphometric data. The clusters merely represent con- centrations of specimens, which theoretically can over- lap at the margins. Since the present work is prelimi- nary, it is possible that some of the marginal specimens may be misassigned to a cluster. The important con- tribution of this study lies in clearly identifying the Text-figure 7.—Scanning electron microscope photographs of modern Porites astreoides from different reef habitats near Discovery Bay, Jamaica. All photos, x40. (A) SUI 49915, sand channel (20 m), (B) SUI 47086, lagoon (16 m), (C) SUI 49935, backreef (1 m), (D) SUI 49925, nearshore patch reef (4 m). Study shows that: the wall reticulum is best developed in the lagoon and thinnest in the patch reef, corallite diameter is largest in the forereef and smallest in the patch reef, corallite diameter is largest in the backreef and smallest in the patch reef, calice elevation is deepest in the lagoon and shallowest in the backreef, thickness of septal structures is greatest in the sand channel and smallest in the lagoon and patch reef. DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 59 number of groups and their centroids, and roughly estimating the variability of each. Within this frame- work, all available primary types from the Caribbean Neogene have been re-evaluated. Also possible rela- tionships with modern Caribbean and Neogene Med- iterranean species have been postulated. MATERIAL The material studied consists of all the specimens of Poritidae (approximately 450 colonies) collected in the Dominican Republic by J. Geister, P. Jung, J. B. Saunders, and other coworkers between 1978 and 1980 (Saunders, Jung, and Biju-Duval, 1986), and is cur- rently deposited at the Naturhistorisches Museum in Basel. These coral collections from the Dominican Re- public are termed “NMB” collections in the following discussion in order to distinguish them from other type and comparative material used in the analyses. First, the Poritidae were separated from the rest of the coral collections and sorted into three genera, Porites, Go- niopora Blainville, 1830, and Alveopora Blainville, 1830. The chief characteristic used to distinguish the three genera was septal number and arrangement (see description of septal structure and arrangement in the following section on characters). Specimens assigned to Porites had 10 to 14 septa, usually consisting of one directive, a ventral triplet, and four lateral pairs. Spec- imens assigned to Goniopora had 18 to 30 septa ar- ranged in three cycles, the first of which consists of five to eight directives. Specimens assigned to Alveopora had weakly-developed septa, not arranged in any def- inite pattern. The specimens within each genus were then divided into groups on the basis of colony shape and corallite diameter. In this process, 10 groups were distinguished in Porites and two in Goniopora. Then, 10 to 15 well-preserved specimens from a wide range of localities were selected from each group. These spec- imens were measured and used in the statistical anal- yses. A total of 120 NMB colonies were selected and Measured. Measurements were also made on 40 type specimens of 22 of the 25 described species of Porites and Go- niopora from the Neogene of the Caribbean region (Table 1). Types of the remaining three described Species, Goniopora calhounensis Weisbord, 1971, Po- rites chipolanum Weisbord, 1971, and Goniopora au- cillana Weisbord, 1973, are currently deposited at Florida State University, Tallahassee, FL, U.S.A., but were not received on loan until after the statistical analyses were completed. In addition to Neogene Ca- ribbean types, specimens of six species of modern Po- rites from the Caribbean region and types of some of the more common Mediterranean Neogene poritid Species were measured (Table 2). Five corallites from various positions within each colony were measured. Previous work (Foster, 1985) has indicated that five is the minimum number of corallites necessary to describe a colony. Since the goal of the present study is essentially exploratory in nature and time was limited, only the minimum number of corallites on the maximum number of colonies were measured. Except where stated, the statistical analyses were performed using colony means of the five cor- allites. To describe and explain the variation within each taxon 1n more detail, subsequent measurement of 10 to 15 corallites per colony as well as simulation of original colony form would be necessary. CHARACTERS The characters analyzed consist of linear measure- ments and counts on 29 calice features (Tables 3, 4; Text-fig. 8). All data are available on computer tape from the author. This emphasis on calices rather than colony form follows the suggestion of Brakel (1976) that corallite (not colony) variation is largely geneti- cally controlled. This approach conforms to the prac- tice of Veron and Pichon (1982) in their recent revision of the modern Poritidae of eastern Australia. Study of calice morphology also represents the most consistent way to analyze fossils for which only fragments of the original colony are available for study. Thin-sections of the Poritidae provide few additional characters be- cause calices are typically shallow and not traceable through the corallum. The characters have been selected to include all di- agnostic features used to distinguish modern poritid species in both the Indo-Pacific and Caribbean regions. They can be grouped into five major categories: (1) corallite size and elevation, (2) septal structure and arrangement, (3) development of the pali, (4) size and structure of the columella tubercle, and (5) size and structure of the coenosteal reticulum. Very little is known ofthe ontogenetic or evolutionary development of these features. Their structure has been described most comprehensively by Bernard (1900, 1903, 1905, 1906). A brief summary is given below: (1) Corallite size and elevation. — As explained above, calices are usually superficial and do not extend deep into the corallum, especially in Porites. They are either circular or polygonal, depending on whether or not the wall reticulum (often termed “coenosteal reticulum") is developed. The calice is bilaterally symmetrical. One measure (an average of measurements made parallel and perpendicular to the line of symmetry), CD, was analyzed to describe corallite size. Four measures, CE, PE, SE, WE, were made to describe overall calice elevation and the elevation of various structures in the calice. Fossa shape and the slope of the septal margins within the calice can be approximated using a com- bination of these measurements. BULLETIN 325 Table 1.—List of Neogene Caribbean poritid types used in statistical analyses. Porites analysis: 1. Porites carrizensis Vaughan, 1917, holotype, USNM 68293, USGS locality 7616, Barrett Canyon, San Diego County, California 2. Goniopora portoricensis Vaughan, 1919, holotype, USNM 325061, USGS locality 3191, Lares, Puerto Rico 3. Goniopora portoricensis Vaughan, 1919, paratype, USNM 325060, USGS locality 3191, Lares, Puerto Rico 4. Goniopora clevei Vaughan, 1919, holotype, PIU УУ110, Anguilla 5. Goniopora clevei Vaughan, 1919, paratype, PIU WI9, An- guilla 6. Goniopora clevei Vaughan, 1919, paratype, USNM 325111, USGS locality 6893, Crocus Bay, Anguilla 7. Goniopora clevei Vaughan, 1919, paratype USNM 523115, USGS locality 6966, Crocus Bay, Anguilla 8. Goniopora clevei Vaughan, 1919, figured specimen, USNM 325116, USGS locality 6016, Empire Quarry, Canal Zone 9. Goniopora cascadensis Vaughan, 1919, holotype, USNM 325072, USGS locality 6020c, Las Cascadas, Canal Zone 10-11. Goniopora cascadensis Vaughan, 1919, paratypes, USNM 335073, USGS locality 6020c, Las Cascadas, Canal Zone 12. Porites baracoaensis Vaughan, 1919, holotype, USNM 325069, USGS locality 3476, Baracoa, Cuba 13. Porites baracoaensis matanzasensis Vaughan, 1919, ho- lotype, USNM 325067a, USGS locality 3461, Matanzas, Cuba 14. Porites baracoaensis matanzasensis Vaughan, 1919, para- type, USNM 3250675, USGS locality 3461, Matanzas, Cuba 15-16. Porites douvillei Vaughan, 1919, original syntypes, USNM 325106 (lectotype designated herein: USNM 325106), USGS locality 6016, Empire Quarry, Canal Zone 17. Porites toulai Vaughan, 1919, holotype, USNM 325105a, USGS locality 6016, Empire Quarry, Canal Zone 18-19. Porites toulai Vaughan, 1919, paratypes, USNM 325105b, USGS locality 6016, Empire Quarry, Canal Zone 20. Porites panamensis Vaughan, 1919, holotype, USNM 325063, USGS locality 6015, Empire Quarry, Canal Zone 21-22. Porites panamensis Vaughan, 1919, paratypes, USNM 325064, USGS locality 6016, Empire Quarry, Canal Zone 23. Porites anguillensis Vaughan, 1919, holotype, PIU W143, Anguilla 24-25. Porites (Synaraea) howei Vaughan, 1919, original syn- types, USNM 325113 (lectotype designated herein: USNM 72890), USGS locality 6016, Empire Quarry, Canal Zone 26—28. Porites (Synaraea) macdonaldi Vaughan, 1919, original syntypes, USNM 325046a (lectotype designated herein: USNM 72376), USGS locality 6016, Empire Quarry, Ca- nal Zone 29. Porites trinitatis Vaughan, in Vaughan and Hoffmeister, 1926, holotype, USNM 353674, USGS locality 8299, Cu- muto Road, Trinidad 30. “Porites floridaprima Bernard” Weisbord, 1973, topo- type, USNM 66220, USGS locality 3286, Tampa, Florida 31. Goniopora ballistensis Weisbord, 1973, paratype, USNM 68314, USGS locality 3286, Tampa, Florida 32. Goniopora matsoni Weisbord, 1973, holotype, USNM 68315, USGS locality 6546, Tampa, Florida Goniopora analysis: 1. Goniopora hilli Vaughan, 1919, holotype, USNM 325058, USGS locality 6016, Empire Quarry, Canal Zone 2. Goniopora panamensis Vaughan, 1919, holotype, USNM 325053, USGS locality 6016, Empire Quarry, Canal Zone 3. Goniopora decaturensis Vaughan, 1919, holotype, USNM 325031, USGS locality 3383, Hales Landing, Georgia 4. Goniopora jacobiana Vaughan, 1919, holotype, USNM 325077, USGS locality 3446, La Cruz, Cuba 5. Goniopora jacobiana Vaughan, 1919, paratype, USNM 325082, USGS locality 6775, White Springs, Florida 6. Goniopora imperatoris Vaughan, 1919, holotype, USNM 325049, USGS locality 6016, Empire Quarry, Canal Zone 7. Goniopora canalis Vaughan, 1919, original syntype des- ignated as lectotype herein, USNM 72891, USGS locality 6016, Empire Quarry, Canal Zone 8. Goniopora tampaensis Weisbord, 1973, holotype, USNM 68317, USGS locality 2084, Tampa, Florida (2) Septal structure and arrangement. — The septa are composed of vertical trabeculae joined by hori- zontal septal junctions and synapticulae. These junc- tions outline the pores, diagnostic of the poritids. A granule (herein termed “denticle”) is formed on the surface of each trabecula. Two types of measures de- scribe the vertical trabeculae: (a) number of denticles per septum (DS), a measure which can be used to estimate number of trabeculae per septum and (b) sep- tum thickness (T1, T2, T3), which can be used to es- timate the size of respective trabeculae. Similarly, two types of measures describe the synapticulae: (a) num- ber of synapticular rings (SR), and (b) synapticula thickness (SY). The septa are bilaterally arranged in a diagnostic formula described in detail by Bernard (1903, 1905). They are commonly termed “dorsal” (one directive septum), “ventral” (a septal triplet), and “lateral” (four septal pairs) in Porites, and “primary” (one cycle of six directive septa), “secondary” (one cycle of six septa), and “tertiary” (one cycle of twelve minor septa) in Goniopora (see Text-fig. 8 for orientation). The total number of septa per calice (NS) was first counted, and the pattern of septal fusion was noted (F1, F2 in Go- niopora). The ventral triplet was almost always fused in Porites and rarely free. A trident (sensu Bernard, 1905) was never seen. The lengths of each type of septum (L1, L2, L3) were measured in both genera, and any apparent “bifurcations” near the wall (NB) were noted. In the material studied, such bifurcations were usually caused by decreased calcification rather than by actual forking of trabecular rows. They there- fore do not represent incipient septa. (3) Development of the pali.—The pali are not true “рай” [defined in Vaughan and Wells (1 943) and Wells (1956) as separate, discrete structures formed by septal substitution], but are similar to paliform lobes, formed by development of the innermost septal trabeculae DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 61 0210 N u, SE SCHULEN V „ Мм Text-figure 8.—Drawings showing some of the characters measured. (A) Porites: corallite diameter, CD; dorsal septum length, Li; ventral Septum length, L2; lateral septum length, L3; ventral palus thickness, P2; columella tubercle length, C1; columellar synapticular ring diameter, CW; wall thickness, WT. (B) Goniopora: corallite diameter, CD; primary septum length, L1; secondary septum length, L2; tertiary septum length, L3; secondary palus thickness, P2; columella tubercle length, C1; columella tangle width, CW; wall thickness, WT. 62 BULLETIN 325 Table 2.—List of Neogene Mediterranean poritid types and modern Caribbean specimens used in statistical analyses. Neogene Mediterranean: 1. Porites collegniana Michelin, 1842, figured specimen in Chevalier (1961), MHNP #P-324-C, Mérignac, Aquitaine, France 2. Porites incrustans Milne-Edwards and Haime, 1851, figured specimen in Reuss (1872) NHMW 1859-XLV-649b, Forchtenau, Hungary . Porites maigensis Kühn, 1925, holotype, Eggenburg Mu- seum, Maigen near Eggenburg, Austria . Porites pachysepta Chevalier, 1961, holotype, MHNP, Vil- landraut, Aquitaine, France 5. Porites mancietensis Chevalier, 1961, holotype, MHNP, Manciet, Aquitaine, France . Porites lobatosepta Chevalier, 1961, holotype, MHNP, Po- pogna, Livorno, Italy . Porites calabricae Chevalier, 1961, holotype, MHNP, Vibo Valentia, Calabria, Italy 8. Porites leptoclada Reuss, 1872, holotype, NHMW 1872-XIII- 80, Niederleis near Enzefeld, Austria 9. Goniopora globulosa Chevalier, 1961, holotype, MHNP, St. Paul-Les-Dax, Aquitaine, France 10. Goniopora granosa Chevalier, 1961, holotype, MHNP, Peyr- ére de Peyrehorade, Aquitaine, France 11. Goniopora turonensis Chevalier, 1961, holotype, MHNP, Pontlevoy, Touraine, France Modern Caribbean: 1. Porites astreoides Lamarck, 1816, holotype, MHNP, La- marck Collection #149, Mers d’Amérique 2. Porites astreoides Lamarck, 1816, USNM 15521, Dry Tor- tugas دیا A о - 3-4. Porites astreoides Lamarck, 1816, YPM 7077—7078, Laborel Collection, Recife Military Air Base, Brazil 5. Porites astreoides braziliensis Verrill, 1901a, holotype, YPM 4539, Maria, Farinha, Pernambuco, Brazil 6. Porites porites (Pallas, 1766), Senckenburg Museum, Esper Collection #19, locality unknown 7. Porites porites (Pallas, 1766), USNM 72375, Florida Keys, locality C-12 8. Porites clavaria Lamarck, 1816, holotype, MHNP, Lamarck Collection #150, Antilles 9. Porites clavaria Lamarck, 1816, USNM 15860, Dry Tortu- gas 10. Porites furcata Lamarck, 1816, holotype, MHNP, Lamarck Collection #154, Mers d’Amérique 11. Porites furcata var. 2 of Lamarck, 1816, MHNP, Lamarck Collection #155, Mers d’Amérique 12. Porites furcata Lamarck, 1816, USNM 15860, Dry Tortugas 13. Porites divaricata Lesueur, 1821, USNM 72377, locality C-45, east of Pulaski Bay, Florida 14. Porites branneri Rathbun, 1887, holotype, USNM 10961, Parahyba do Norte, Brazil 15. Porites branneri Rathbun, 1887, paratype, USNM 10962, Canderas Reef, Pernambuco, Brazil 16. Porites branneri Rathbun, 1887, YPM 9079, Laborel Col- lection, Abrolhos Archipelago, Brazil 17. Porites branneri Rathbun, 1887, YPM 9080, Laborel Col- lection, Fernando de Novonha Is., Brazil 18. Porites verrilli Rehberg, 1893, holotype, YPM 4539, Albrolhos Reefs, Brazil (Text-fig. 9). For consistency with other previous po- ritid studies, the term “palus” is nevertheless retained. The number of pali (PL) was counted within each cali- ce, and the widths of those corresponding to the three septal types (P1, P2, P3) measured. A synapticular ring connecting all pali within each calice was often found in Porites and the distance across this ring (CW) was measured. (4) Size and structure of the columella tubercle. — In Porites, the columella consists of an isolated tubercle aligned parallel to the dorsoventral axis and supported by the columellar synapticular ring connecting the pali. In Goniopora, the columella consists of a tangle of twisted trabeculae extending from the inner margins of the septa. In Porites, the length (C1) and the width (C2) of the tubercle were measured. In Goniopora, the length (C1) and the width (C2) of an upper lobe of a central trabecula (termed ““tubercle”) were measured, as well as the width of the entire columella tangle (CW). (5) Size and structure of the coenosteal reticulum. — is Text-figure 9.—Serial acetate peels down one corallite of Porites furcata. (A), (B), and (C) were made at regular 0.1-mm intervals from the colony surface, and (D) was made 0.3 mm below (С). In (A), six pali are clearly seen surrounding a thin columella tubercle. In (В), the pali are five in number and begin to blend with the septa. The dorsal palus has already merged. In (C), the pali are no longer visible and the corallite outline begins to disappear into an irregular (faintly radial) meshwork of septal trabeculae. In (D), the corallite has disappeared altogether, and the skeletal meshwork appears structureless and more heavily calcified. DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 63 As described in detail by Bernard (1905), the wall or theca consists of one or more rows of vertical trabec- ulae united by synapticulae. When more than one row is involved, the trabecular complex is usually termed “wall reticulum" or even “‘coenosteal reticulum". The reticulum is not true coenosteum, however, but is formed by addition of discrete rows of intervening mu- ral trabeculae and not by accretion or deposition of structurally-different skeletal material. This coenosteal reticulum is the only skeletal material that separates corallites. It may form bulges or ridges across the ca- lical surface. Three measures describe the amount of coenosteal reticulum separating calices: corallite spac- ing (CS), number of rows of mural trabeculae (WR), and wall thickness (WT). One additional measure (to- tal number of mural denticles, DW) describes the over- all texture of this reticulum. STATISTICAL PROCEDURES Species were distinguished within each of the two common genera, Porites and Goniopora, using multi- variate statistical analyses similar to those outlined in Foster (1984). First, the means for each colony in the NMB collections were assigned to preliminary groups within each genus using cluster analysis (BMDP pro- gram 2M, Dixon, 1981). All measured characters were used in the analyses. Eight groups were formed in Po- rites and four in Goniopora. Next, these groups were re-analyzed using a series of stepwise discriminant analyses (BMDP program 7M, Dixon, 1981). In this process, analyses were also run using data for individual corallites, and group assign- ments for colonies were changed so that all corallites within each colony belonged to the same group. In cases showing extensive overlap between groups, the overlapping groups were analyzed separately. If the groups still showed considerable overlap, the groups were combined and then re-evaluated together with all groups within the genus. Group assignments for col- onies lying at cluster margins were further modified by trial and error to obtain the highest percentage of cor- rectly-classified corallites. Because of poor preserva- tion of some of the type specimens, a few characters (primarily related to thickness) were dropped from the final analyses. All measured characters except DS, SR, SD, and SY were used in the final Porites analysis. All characters except F1, F2, DS, T1, T2, T3, SD, and SY were used in the final Goniopora analysis. The final results yielded five groups among the NMB specimens in Porites (Text-fig. 10a) and three in Goniopora (Text- fig. 11a). These groups represent the species described in this study. Finally, using the NMB species groups and adding the measured types from Table 1, two further discrim- inant analyses (BMDP program 7M, Dixon, 1981) were performed: (1) the analyses were rerun leaving the types unclassified, (2) the analyses were rerun so that each NMB group and group of types were classified sepa- rately. The results (Text-figs. 10b, 11b) revealed a very complicated relationship between the types and the Table 5.— Weighting of characters in the all Porites canonical discriminant analysis. Total-sample correlations between the canonical variables and the original variables (COR), and standardized canonical coefficients (SCC). Only values with high magnitudes are given. The four canonical variables are labelled CV 1—4. Abbreviations for characters are explained in Table 3. u CVI СУ2 СУЗ СУ4 Original variable COR SCC COR SCC COR SCC COR SCC CD - 0.87 —0.422 — —0.51 —0.340 = cs 0.764 = = 10011 0.288 1.10* = 0.81 L1 = = —0.610* —1.19 = = = = L2 = А2. —0.492 - 0.71 E E L3 = = —0.574 - = = 0.58 NN = = = = = 0.467 0.73 NB 0.616 = > 0.263 0.54 = = DW —0.538 = —0.457 — ر‎ — - - PL —0.522 = = = —0.65 E = WR 0.900* 1.47* ES —1.75* = = = = TI = = = = = —0.487 = T2 = = = 0.316 0.84 —0.536* = T3 = = = = = —0.458 = P2 0.530 - - = = = Cl 0.648 = - = —0.43 = = C2 0.578 E = = —0.50 = = Cw 0.667 = = 0.96 0.264 = 0.474 —1.50* WT 0.905* E = 1.90* = = = s SE = =0.72 = =) Sie —0.65 нь —0.75 WE 0.607 0.89 E 0) sly —0.69 = 0.92 * most important variables. 64 BULLETIN 325 B4 b с 4 А 4-4 м] 9 N I с А Е 09-4 У А я І А B a L 4 E -4- 2 4 * -B €———————Á— M — -8 -4 0 4 8 CANONICAL VAAIABLE 1 8-і ANE ET | en A 4- N 4 0 N 1 c A L 4 04 v d A R 1 А B ise | E -44 Йй - Р 2 4 -8 рр Т -8 -4 0 4 8 B CANONICAL VARIABLE 1 Text-figure 10.— Porites. Canonical discriminant analysis of the NMB collections. (A) Plot of scores on the first two canonical variables showing polygons outlining the range of variation between colonies in the five species groups defined by the statistical analyses. The points represent means for each colony. 1 — P. portoricensis, 2 — P. baracoaensis, 3 — P. waylandi, 4 — P. convivatoris, 5 — P. macdonaldi. (B) Plot of scores showing polygons around the NMB species groups [as given in (A)] and points representing the colony means of the measured types. A = type 23, B = types 12-14, C = types 9-11, D = types 15-16, F = type 30, H = types 24-25, L = type 31, M = types 26-28, N = types 20-22, P = types 2-3, R = type 29, S = type 32, T = types 17-19, V = types 4-8, Z = type 1. The numbers for types refer to specimens listed in Table 1. DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 65 г»он2 0 2 > О № 1 si a EN E eb — -. mb E nruo>HD>< N 1 سے‎ Rr Me Nu м Nm S № N A a сақ —— ج‎ F. PAF ЧУ. a ы u 04 S % Ы / = 2 q Y 2 -4- -Б- Т T 1 T T T T -6 ze B 4 6 8 10 CANONICAL VARIABLE 1 A 64 4-4 T»OHZOZ»O го 1 04 У А R I A 24 B ІШ H Е Р 7 2 -4- = T T T T T T T -8 -4 2 4 6 8 10 CANONICAL VARIABLE 1 Text-figure 1 1.—Goniopora. Canonical discriminant analysis of the NMB collections. (A) Plot of scores on the first two canonical variables showing polygons outlining the range of variation between colonies in the three species groups defined by the statistical analyses. The points represent individual corallites. 1 = G. hilli, 2 = G. imperatoris, 3 = G. calhounensis. (В) Plot of scores showing polygons around the NMB Species groups [as given in (A)] and points representing the colony means of the measured types. C = type 7, D = type 3, H = type 1,1 = type 6, J = types 4-5, P = type 2, T = type 8. The numbers for types refer to specimens listed in Table 1. 66 BULLETIN 325 FREQUENCY 14 13 0 CANONICAL VARIABLE 1 BA rpnHmHzoz>Nn mnrop pHD>< -44 го -84 T T T T T T T BER 5 3 Y Y Y Y Y T T T T T т T y T T T T T T T T T 7T LE T T MEAT -8 -4 0 4 8 CANONICAL VARIABLE 1 Text-figure 12.— Porites subgroups. Canonical discriminant analysis of the NMB collections. (A) Groups 2 and 4, P. baracoaensis and P. convivatoris. Histogram giving frequencies of colony means for scores on the canonical variable. Left angle stripes - NMB P. baracoaensis; right angle stripes = NMB P. convivatoris; solid = types 12-14; small crosshatch = types 17-19; large crosshatch — types 15-16. The numbers for types refer to specimens listed in Table 1. (B) Groups 1, 3, and 5; P. portoricensis, P. waylandi, and P. macdonaldi. Plot of scores on the first two canonical variables showing polygons outlining the range of variation between colonies in the NMB Species groups. The points represent means for each colony. 1 = NMB P. portoricensis, 3 = NMB P. waylandi, 5 = P. macdonaldi, A = type 23, C = types 9-11, F = type 30, H = types 24—25, L = type 31, M = types 26-28, N = types 20-22, P = types 2-3, S = type 32, V = types 4-8. The numbers for types refer to specimens listed in Table 1. DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 67 NMB species groups. To interpret this relationship, the Porites were divided into two subsets, the first com- posed of NMB species groups 2 (P. baracoaensis Vaughan, 1919) and 4 (P. convivatoris, n. sp.) and the second encompassing NMB species groups 1 [P. por- toricensis (Vaughan, 1919)], 3 (Р. waylandi, nom. nov.), Table 6.— Weighting of characters in the all Goniopora canonical discriminant analysis. Total-sample correlations between the ca- nonical variables and the original variables (COR), and standardized canonical coefficients (SCC). Only values with high magnitudes are given. The canonical variables are labelled CV1-2. Abbreviations for characters are explained in Table 4. and 5 (P. macdonaldi Vaughan, 1919). Each subset was Original 144 ME reanalyzed together with only those types that origi- variable СОК SCC COR SCC nally lay near the constituent NMB species groups. The CD 0.918* 1.04* = = results are shown in Text-figure 12. Similarly in Go- cs Many 0.54 - un niopora, NMB species groups 1 (G. hilli Vaughan, 1919) а EE = T 030 ; : 12 0.907* — — 0.93* and 2 (С. imperatoris Vaughan, 1919) have been ге- 13 0.910* 0.91* A жы analyzed alone with all the Goniopora types (Text-fig. DW 0.657 S = = 13). These analyses form the basis of: (1) the syno- PL = = —0.510* —0.59 nymies and the distribution patterns given in the formal SR 0.520 ж 0.349 0.65 Systematic descriptions and (2) the brief discussions 52 4 ЩЕ 658 mend le uw given on biostratigraphy and paleoecology. The results CW 0.633 Үр —0.332 ad shown on Text-figures 10 through 13, Tables 5 and 6, SY 0.652 0.50 аш 366 a and Appendix Па—а are based on the results of the SAS WT 0.596 - -0.339 -0.64 CANDISC procedure (SAS Institute, 1982) run at the SE is = 0.452 0.72 * most important variables. FREQUENCY 20 + | N | S N 15 - N N — NS SS АА 2 | NNN A 2 2 \ \ S EV de SS МҸ МҸ МҸ Қ Ж Ж 2 2 -4 -3 -2 T 0 1 2 3 4 5 6 CANONICAL VARIABLE 1 Text-figure 13.—Goniopora subgroup. Canonical discriminant analysis of the NMB collections. Histogram giving frequencies of colony Means of scores on the canonical variable. Left angle stripes = NMB G. imperatoris; right angle stripes = NMB G. hilli, small left angle stripes = type 7; small right angle stripes = type 1; solid = type 8; small crosshatch = type 3; medium crosshatch = type 6; large crosshatch = types 4— 5. The numbers for types refer to specimens listed in Table 1. 68 University of Iowa. In all Text-figures, Tables, and Appendices, the data used in Porites represent colony means, whereas in Goniopora, they represent individ- ual corallites. Means and standard deviations of all single char- acters are given for each species in Appendix Ia, b. Eight characters revealing highly-significant differences between species are plotted for Porites in Text-figure 14 and for Goniopora in Text-figure 15. Intraspecific variation was found to be roughly equivalent in the calculated canonical variables for all species in each genus using Box’s test (Miller, 1968). RESULTS AND INTERPRETATIONS As described in the previous section, five separate canonical discriminant analyses were performed to dis- tinguish species and relate them to type material: (1) all five groups of Porites, (2) groups 2 and 4 of Porites, (3) groups 1, 3, and 5 of Porites, (4) all three groups of Goniopora, and (5) groups 1 and 2 of Goniopora. The results of each analysis are as follows: BULLETIN 325 (1) All five groups of Porites: 1, P. portoricensis; 2, P. baracoaensis; 3, P. waylandi; 4, P. convivatoris; 5, P. macdonaldi. — Five groups were distinguished in this analysis, with 90.2 percent of all colonies classified correctly (Text-fig. 10a). 80 percent were correctly clas- sified 1n groups 1 and 4, 93.3 percent in groups 3 and 5, and 100 percent in group 2. Four canonical variables (CV 1-CV4) were calculated, with CV1 accounting for 77.7 percent of the variation, CV2 for 11.4 percent, CV3 for 6.7 percent, and CV4 for 4.2 percent. The initial stepwise analysis (computed using the program BMDP-7M) showed that the groups could be distin- guished using a minimum of seven characters (in de- creasing importance: WE, L1, CW, NB, WR, NN, and SE). However, 20 of the 25 measured characters were important in computing the character combinations that best distinguished the five species in the total ca- nonical analysis (computed using the SAS CANDISC procedure) (Table 5). The first canonical variable (CV 1) distinguished three major groups: group 2, group 4, and a combination of groups 1, 3, and 5. It weighted POR(N=30) сео ӨЕ POR(N=30) тер POR(N=30) ae BAR(N=32) == BAR(N=32) Ser BAR(N=32) =e way (N=15) = WAY (N=15) Ж; WAY (N=) 74 CON(N=10) po — CON(N=10) xe CON(N=10) 29 MAC(N=15) Bern ae MAC(N=15) 229 MAC(N=15) ne таа T T T T T » Ц Т | 1 1.5 2 0.2 0.4 0.6 0.8 4 6 A. CORALLITE DIAMETER (MM) B. DORSAL SEPTUM LENGTH (MM) С. NUMBER OF PALI POR(N=30) а= POR(N=30) как POR(N=30) Б EEE BAR(N=32) м BAR(N=32) = BAR(N=32) == WAY (N=15) == WAY (N=15) “Sz WAY (N=15) 2%- CON(N=10) e CON(N=10) T9 CON(N-10 | —9— MAC(N=15) AE MAC(N=15) enas MAC(N=15) Eb i 0 2 4 6 0.05 010 045 0.20 0.2 04 06 оз D. ROWS OF MURAL TRABECULAE E, VENTRAL SEPTUM THICKNESS (MM) F COLUMELLA SYN. RING DIAMETER (MM) POR(N=30) =. POR(N=30) ERE BAR(N=32) e BAR(N=32) EJ WAY (N=15) pee WAY (N=15) TE CON(N=10) -% CON(N-10) m 4 MAC(N=15) та MAC(N=15) ar 0 0.5 1 07 0.5 1 С. WALL THICKNESS (MM) H. CALICE ELEVATION (MM) Text-figure 14.— Means and standard deviations for eight characters in the five Porites species. The midpoint ofeach horizontal line represents the mean, and the length of the line on either side of the midpoint is one standard deviation. Analysis of variance (or Welch’s statistic when variances are unequal) shows that the species are significantly different in each case. Levene’s test for equal variances shows that variances are equal only in dorsal septum length. POR = species 1, P. portoricensis, BAR = species 2, P. baracoaensis; WAY = species 3, P. waylandi, CON = species 4, Р. convivatoris, MAC = species 5, Р. macdonaldi. “М” = number of colonies measured. (A) corallite diameter, CD, (В) dorsal septum length, L1, (C) number of pali, PL, (D) number of rows of mural trabeculae, WR, (E) ventral septum thickness, T2, (F) columellar synapticular ring diameter, CW, (G) wall thickness, WT, (H) total calice elevation, WE. DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 69 number of rows of mural trabeculae (WR) most heavily and is strongly correlated with the number of rows of mural trabeculae (WR) and wall thickness (WT). The second canonical variable (CV2) distinguished be- tween groups 1, 3, and 5. It again weighted the number of rows of mural trabeculae (WR) heavily but is best correlated with the inverse of the lengths of the three septal types (L1, L2, L3). Various characters are heavi- ly weighted on canonical variables 3 and 4 (CV3, CV4), with corallite spacing (CS) most important in CV3 and total columella width (CW) most important in CV4. Two measures of calice elevation (SE, WE) are in- versely correlated with CV3, and septal thickness (T1, T2) is inversely correlated with CV4. (2) Groups 2 (P. baracoaensis) and 4 (P. convivatoris) of Porites. — Two groups were distinguished in this analysis, with 100 percent of all colonies correctly clas- sified (Text-fig. 12a). The canonical variable calculated explained 100 percent of the variation. Nine of the 25 characters were important in the total canonical anal- ysis (Appendix Па). Columella tubercle length (C1) was most heavily weighted in the canonical variable, and calice elevation (WE) is most strongly correlated. The canonical variable appears most similar to CV3 of the first analysis. Study of the position of the types on the histogram for the canonical variable shows that types for three described species belong to group 2, whereas no types belong to group 4. (3) Groups 1 (Р. portoricensis), 3 (Р. waylandi), and 5 (P. macdonaldi) of Porites. —Three groups were dis- tinguished in this analysis, with 85 percent of all col- onies classified correctly (Text-fig. 12b). 85.7 percent were correctly classified in groups 3 and 5, and 83.3 percent in group 1. Two canonical variables were cal- culated, which explained 74.7 percent and 25.3 percent of the variation respectively. Fifteen of the 25 char- acters were important in the total canonical analysis (Appendix IIb). Septum length (L1) was most heavily weighted and most strongly correlated on the first ca- nonical variable (CV 1), which appears most similar to HIL(N=6) 9; HIL(N=6) HIL(N=6) EZ IL IMP(N=10) a IMP(N=10) DE IMP(N=10) ——— CAL (N=2) Sora CAL (N=2) eae CAL (N=2) es | eee T T 1 Fr T 1 [ T 1 A 0 2 4 6 0.5 1.5 2 20 25 30 * CORALLITE DIAMETER (MM) B. DORSAL SEPTUM LENGTH (MM) C. TOTAL NUMBER OF SEPTA HIL(N=6) En We HIL(N=6) se HIL(N=6) -%- IMP(N=10) a IMP(N=10) = IMP(N=10) ^9 CAL (N=2) "hr CAL (N=2) ж CAL (N=2) EL r T T 1 r 1 r T T 1 8 10 12 м 0.5 1.5 0.05 010 0.15 020 D, NUMBER OF PALI Gm TOTAL COLUMELLA WIDTH (MM) E SYNAPTICULA THICKNESS (MM) HIL(N=6) Ta HIL(N=6) Sa IMP(N=10) er IMP(N=10) ЕНЕ CAL (№2) > CAL (N=2) = r T T Te 1 0.4 0.6 0.8 1 12 0.2 04 0.6 с. WALL THICKNESS (MM) 0.0 Н. SEPTA ELEVATION (MM Text-figure 15. — Means and standard deviations for eight characters in the three Goniopora species. The midpoint of each horizontal line Tepresents the mean, and the length of the line on either side of the midpoint is one standard deviation. Analysis of variance (or Welch's Statistic when variances are unequal) shows that the species are significantly different in each case. Levene's test for equal variances shows that variances are unequal only in corallite diameter, dorsal septum length, and synapticula thickness. HIL = species 1, G. hilli; IMP = species 2, G. imperatoris, CAL = species 3, G. calhounensis. *N" = number of colonies measured. (A) corallite diameter, CD, (B) primary septum length, L1, (C) total number of septa, NS, (D) number of pali, PL, (E) columella tangle thickness, CW, (F) synapticula thickness, SY, (G) wall thickness, WT, (H) septa elevation, SE. 70 BULLETIN 325 CV2 in the first analysis. Total columella width (CW) was most heavily weighted on CV2, which is best cor- related with the number of neighboring corallites (NN), the inverse of septum thickness (T2), and the inverse of total columella width (CW). CV2 appears most sim- ilar to CV4 of the first analysis. Study of the distances of types from these three group centroids shows that: (1) two described species belong to group 1, (2) four described species belong to group 3, (3) one described species belongs to group 5, and (4) the positions of two described species lie intermediately between groups 1 and 3. Further subjective examination of variation in overall colony shape within these three species shows that the two intermediate described species most likely belong to group 1. This information together with the poor preservation of types for the two additional de- scribed species that initially appeared to belong to group 3 suggests that they may instead belong to group 1. (4) All three groups of Goniopora: 1, G. hilli; 2, G. imperatoris; 3, G. calhounensis.— Three groups were distinguished in this analysis, with 91.1 percent of all corallites correctly classified (Text-fig. 11a). 92 percent were correctly classified in group 2, and 90 percent were correctly classified in groups 1 and 3. Two ca- nonical variables (CV1, CV2) were calculated, with the first accounting for 93.6 percent of the variation and the second for only 6.4 percent. The stepwise discrim- inant analysis showed that only four (in descending order: CD, L3, PL, SR) of the 21 characters were need- ed to distinguish between the three groups. However, in the total canonical analysis, 14 of the 21 measured characters were important (Table 6). The first canon- ical variable (CV1) distinguished the three groups, al- though overlap between groups | and 2 still persisted. It weighted corallite diameter (CD) and tertiary septum length (L3) most heavily and is very strongly correlated with corallite diameter (CD) and septum length (L1, 1.2, L3). The second canonical variable (CV2) pri- marily distinguished groups 1 and 2. It also weighted septum length (L1, L2) most heavily but is inversely correlated with number of pali (PL). Study of the dis- tances of the types from these three group centroids (Text-fig. 11b) shows that no types belong to group 3, and types for seven described species lie near groups 1 and 2. (5) Groups 1 (G. hilli) and 2 (С. imperatoris) of Go- niopora.— Two groups were distinguished in the anal- ysis, with 95 percent of all corallites correctly classified (Text-fig. 13). 93.3 percent were correctly classified in group 1, and 96.0 percent in group 2. The one canonical variable calculated explained 100 percent of the vari- ation. Nine of the 21 characters were important in the total canonical analysis (Appendix IIc). Corallite di- ameter (CD) was most heavily weighted and strongly correlated with the canonical variable. Study of the position of the types on the histogram calculated for the canonical variable shows that types for four de- scribed species belong to group 1, the type of one de- scribed species belongs to group 2, and the type for one described species is intermediate between groups 1 and 2. One additional canonical discriminant anal- ysis was performed classifying three Dominican Re- public groups and all seven described species as 10 separate groups. The results of the analysis clearly in- dicated that the intermediate belonged to group 2. In summary, the results for Porites show that: (1) Five slightly overlapping groups exist. The most overlap occurs in the species having an extensive coe- nosteal reticulum. The overlap is greater than that found in early Miocene species of Montastraea Blainville, 1830 (see Foster, 1984). (2) Almost all the measured characters are important for distinguishing the five groups in the analysis. Four major character complexes were involved in this dis- tinction: (a) development of the wall reticulum; (b) corallite size; (c) calice elevation; and (d) septal and columellar thickness. The pali do not appear to play as important a role in species discrimination as they do in modern Pacific poritids (see Veron and Pichon, 1982). (3) Analyses of type material for 15 described species show that three described species belong to group 2 (P. baracoaensis), possibly six to group 1 (P. portori- censis), two to group 3 (P. waylandi), and one to group 5 (P. macdonaldi). This results in the total synony- mization of eight described species with four previ- ously described species. (4) One new species (group 4, P. convivatoris) is rec- ognized. In summary, the results for Goniopora show that: (1) Three groups exist. The two more abundant groups overlap slightly. (2) Two-thirds of the measured characters are im- portant in distinguishing the groups. Two major char- acter complexes were involved in this distinction: (a) corallite size; and (b) number of pali (and to a lesser degree, calice elevation). (3) Analyses of type material for seven previously described species show that four described species be- long to group 1 (С. hilli), two to group 2 (С. impera- toris), and none to group 3 (G. calhounensis). At the end of the project, however, one type was found that could subjectively be assigned to group 3. This results in the total synonymization of five species with three previously-recognized species. GENERAL COMPARISONS WITH OTHER FAUNAS The results have been statistically compared with DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 71 two separate faunas to obtain further information on the spatial and temporal distribution of each species cluster. These two faunas are: (1) the Miocene Medi- terranean poritids described by Chevalier (1961) and (2) described species of modern Caribbean poritids. The material studied is listed in Table 2. (1) Miocene Mediterranean poritids. — Frost (1977c) has suggested that greater than 50 percent of the Caribbean reef-coral species in the Oligocene also occurred in the Mediterranean region. This overlap occurs especially in the suborder Faviina of the clas- sification system of Vaughan and Wells (1943), a sub- order different from the one under consideration (i.e., the family Poritidae belongs to the suborder Fungiina); however, Frost does report high similarities between three Oligocene species of Goniopora and one Oligo- cene species of A/veopora in the two regions. In con- trast, little overlap is reported between species of Po- rites. No such general comparisons have been made between Caribbean and Mediterranean reef-corals for the Miocene. In the present study, types of 11 Miocene Mediterranean poritid species (three of the five spec- imens of Goniopora and all eight specimens of Porites) described by Chevalier (1961) (Table 2) were mea- sured, and their canonical scores were calculated using the canonical variables of the analyses of the five species of Porites and three species of Goniopora described in the previous section. The results are given in Text- figure 16. In Porites (Text-fig. 16a), the Mediterranean species appear different from the Caribbean species. Only one Species, Р. maigensis Kühn, 1925, has a high CV1 value signifying a well-developed wall reticulum, which is characteristic of Dominican Republic groups 1, 3, and 5 (P. portoricensis, P. waylandi, and P. macdon- aldi, respectively). Similarly, all eight Mediterranean Species have high values for CV2 indicating small-sized Calices. One Mediterranean species, P. mancietensis Chevalier, 1961, is shown to fall within the species 2 Polygon (P. baracoaensis), and three Mediterranean Species, Р. lobatosepta Chevalier, 1961, Р. incrustans Milne-Edwards and Haime, 1851, Р. leptoclada Reuss, 1872, fall within the species 4 polygon (Р. convivatoris). Further study of colony shape (see “Comparison” sec- tions following under Systematic Paleontology), how- ever, indicates that the Caribbean and Mediterranean Species are not synonymous. In Goniopora (Text-fig. 16b), only one Mediterra- nean species appears similar to any Caribbean species (.e., species 1, G. hilli). The corallite diameters of the remaining four species described by Chevalier (1961) are considerably larger than the Caribbean material. The one similar Mediterranean species (G. turonensis Chevalier, 1961), however, again has a distinctly dif- ferent colony shape; therefore, the Caribbean and Med- iterranean species are probably not synonymous. In summary, these analyses suggest that: 1. No Porites or Goniopora species probably co- occur in the Mediterranean and Caribbean regions during the Miocene. 2. The Mediterranean Porites generally have smaller corallites with less-developed wall reticula than the Caribbean Porites. 3. The Mediterranean Goniopora generally have larger corallites than the Caribbean Goniopora. The differences between the Caribbean and Medi- terranean poritids are more likely historical than adap- tive in origin, since the two genera probably assumed similar ecologic roles in the two regions. (2) Modern Caribbean poritids. —Only one of the three genera in the present study, Porites, is represented in the modern Caribbean. Types of six described mod- ern Caribbean species [assuming P. porites (Pallas, 1766) = P. clavaria Lamarck, 1816] and the five Do- minican Republic species groups were analyzed by ca- nonical discriminant analysis using a total of 11 groups. The results are shown on Text-figure 17, and relative loadings for characters are given in Appendix IId. 79.3 percent of the colonies were correctly classified in this analysis. Seven canonical variables were calculated and the first canonical variable (CV 1) explains 58.1 percent of the variation, CV2 19.8 percent, CV3 10.6 percent, CV4 4.8 percent, CV5 4.4 percent, CV6 2.0 percent, and CV7 0.2 percent. Of the two variables shown in Text-figure 17, wall thickness and number of rows of mural trabeculae are most important in CV1, and ca- lice size and the inverse of calice elevation are most important in CV2. Three significant interpretations can be made from these results: (1) Modern Caribbean Porites are distinctly different from the five Neogene Dominican Republic species groups. Overlap occurs only with the species 4 polygon, which represents P. convivatoris, n. sp., an unusual species that forms small ellipsoidal colonies with an axial tube. Therefore, during the late Pliocene or Pleis- tocene, all five ofthe Dominican Republic species must have become extinct, and the modern Porites radiated to replace them ecologically. This hypothesis is sup- ported by the occurrence of four modern species of Porites in the upper Pliocene/lower Pleistocene Ca- loosahatchee Formation of Florida and the absence there of all Neogene poritid species described herein (Weisbord, 1974). This evolutionary pattern in the poritids contrasts with that observed in Montastraea (Foster, in preparation), in which the two modern species are both represented in the Neogene Domini- can Republic material. 72 BULLETIN 325 № no Hi>< ГРО">О?РО о лс. қ ub. ка ЗЫ Me ж «с ЕР N / у: - o z = ө a -8- 1 D 1 ومومو‎ q qíáA A | -В -4 0 4 8 A CANONICAL VARIABLE 1 6-4 4-4 с А N 0 N 24 T 2 с А L 0- У А R 1 1 A 2 B C 3 E 2 2 -4- -6 Т Т T ee T тар -6 -4 2 4 6 8 10 B CANONICAL VARIABLE 1 Text-figure 16.—Classification of Neogene Mediterranean types using the canonical discriminant analyses of the NMB collections. (A) Porites. Plot of scores showing polygons around the NMB species (as given in Text-fig. 10a) and points representing the colony means of measured types. B = type 7, C = type 1, I = type 2, M = type 3, N = type 5, P = type 4, T = type 8. The numbers for types refer to specimens listed in Table 2. (B) Goniopora. Plot of scores showing polygons around the NMB species (as given in Text-fig. 11a) and points representing the colony means of measured types. T = type 11. Types 9 and 10 lie off the graph. The numbers for types refer to specimens listed in Ta- ble 2. DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 73 (2) The modern species have intermediate to high values for CV1 indicating that their coenosteal retic- ulum is generally well-developed and similar to that in species 4 (P. convivatoris) and the species complex 1/3/5 (P. portoricensis, P. waylandi, and P. macdon- aldi, respectively). The highest values for CV1 occur in modern specimens from Brazil. (3) The modern Caribbean species have significantly low values for CV2 indicating that they have smaller, more elevated calices than the fossil forms. Since the fossil material is exquisitely preserved, these results could not be caused by preservation. Further study of additional type material shows that the 1/3/5 species complex (P. portoricensis, P. waylan- di, P. macdonaldi, respectively) depicted in Text-figure 10a and 17 extends into the late Oligocene, and that species 2 (P. baracoaensis) and 4 (P. convivatoris) orig- inate during the Miocene. Apparently, the modern P. astreoides Lamarck, 1816, now replaces the older com- plex, which has a generally similar growth form and wall structure. The modern branching species (P. po- rites, P. furcata Lamarck, 1816, and P. divaricata Le- sueur, 1821) all have a more open wall structure than their fossil counterpart, species 2 (P. baracoaensis). Because of their high abundance and diversity, com- parison with modern Indo-Pacific poritids is beyond the scope of the present investigation. However, it should be noted that, on preliminary examination, these fossil poritids from the Dominican Republic do not resemble any of the modern eastern Pacific poritids reported by Durham (1947). SYSTEMATIC PALEONTOLOGY INTRODUCTION The formal systematic descriptions in this section are based strictly on the results of the statistical anal- yses given in the previous section. Unless otherwise indicated, the synonymies include only material ac- tually measured and statistically analyzed. A list sum- marizing these synonymies is given in Table 7. Fol- lowing Jung (1986), the term “diagnosis” is used only BH ] 4 буша A 44 - о 7] N m 4 ECC A] бек 07 м m R J I - А =ч gu ЕЗ Е -4- Beni (у) А er T T Dun ap X T y EHER eae Re -B -4 0 4 8 CANONICAL VARIABLE 4 Text-figure 17.— Porites. Canonical discriminant analysis of the NMB fossil collections (numbered) and six modern Caribbean species (lettered). Plot of scores on the first two canonical variables showing polygons outlining the range of variation between colonies in the 11 Species groups. The points represent individual colonies. 1 = P. portoricensis, 2 = P. baracoaensis, 3 = P. waylandi, 4 = P. convivatoris, 5 = P. macdonaldi, A = P. astreoides types 1-5, Р. branneri types 14-17, D = Р. divaricata type 13, F = Р. furcata types 10-12, Р = Р. porites types 6-9, V = P. verrilli type 18. The numbers for types refer to specimens listed in Table 2. 74 BULLETIN 325 to describe higher categories such as genera and sub- genera, whereas the term “description” is reserved for complete descriptions of species, which, as discussed in the previous section, herein represent merely mor- phologic clusters of specimens. Lectotypes have been selected for all studied species whose primary types consist of syntypes or ““cotypes”. The terminology used in the “Description” sections is defined and described in the previous section entitled “Characters”. It follows the usage of Bernard (1903, 1905), Vaughan and Wells (1943), and Wells (1956). The abbreviations used in the measurements sections are explained in Tables 3 and 4. These measurements are identical to those used in the statistical analyses. Two additional characters, CH and CT, refer respec- tively to the colony height (in mm) and thickness (in mm). The “Materials” sections give an approximate es- timate of the amount of material in the collection of Saunders, Jung, and Biju-Duval (1986) and the num- ber of specimens in this collection that were statisti- cally analyzed. Species definitions are based on all of these specimens as well as on other specimens listed in the synonymies. The expression “box of fragments" is used to indicate one colony whose fragments actually fit together so that the colony could be reconstructed. Sometimes different boxes of fragments appear to be- long to the same colony; however, the fragments do not fit back together. Because of this problem, esti- mates of possible number of colonies are also given. The “Remarks” sections explain the synonymies only. Separate sections entitled “Variability” describe the variation within each cluster. Sections entitled “Occurrence” give detailed geographic and strati- graphic information within the studied areas of the Dominican Republic, whereas those entitled “Distri- bution” give general information on all known occur- rences throughout the world. Unless otherwise stated, the locality numbers belong to the Naturhistorisches Museum Basel (NMB) and the material is deposited at the NMB. Assignment of formation names to in- dividual localities is based on Saunders, Jung, and Biju- Duval (1986) (especially text-figs. 4, 6, 15, and 16), except along Rio Mao where formation names are used as listed by Maury, 1919. Catalogue numbers have been assigned only to figured specimens. A unique number was given to each colony. Ages used in the “Distribution” sections are based on the following publications: Anguilla, Bold (1970); Antigua, Bold (1966); Cuba, Bold (1975); Jamaica, Zans et al. (1963); Puerto Rico, Frost et al. (1983); Chiapas, Mexico, Frost and Langenheim (1974); Florida and Georgia, Puri and Vernon (1964); Panama, Woodring (1957, 1964); Trin- idad, J. B. Saunders (oral commun., 1985). Table 7.—List of all valid poritid species described from the Mio- cene through Lower Pliocene of the Caribbean region, showing their current taxonomic status. Alveopora fenestrata (Dana) of Duncan, 1863 = P. portoricensis А. tampae Weisbord, 1973 Goniopora aucillana Weisbord, 1973 = G. hilli ballistensis Weisbord, 1973 = P. portoricensis calhounensis Weisbord, 1971 canalis Vaughan, 1919 = G. hilli cascadensis Vaughan, 1919 = Р. portoricensis clevei Vaughan, 1919 = P. portoricensis decaturensis Vaughan, 1919 = ?G. imperatoris hilli Vaughan, 1919 imperatoris Vaughan, 1919 G. jacobiana Vaughan, 1919 = G. hilli G. matsoni Weisbord, 1973 = P. portoricensis G. panamensis Vaughan, 1919 G. tampaensis Weisbord, 1973 = G. hilli Porites anguillensis Vaughan, 1919 astreoides Lamarck of Coryell and Ohlsen, 1929 = P. waylandi baracoaensis Vaughan, 1919 carrizensis Vaughan, 1917 chipolanum Weisbord, 1971 collegniana Michelin of Duncan, 1863 = P. waylandi convivatoris, n. Sp. P. douvillei Vaughan, 1919 — ?P. baracoaensis * P. floridaprima Bernard" Weisbord, 1973 — P. waylandi P. howei Vaughan, 1919 — ?P. portoricensis P. macdonaldi Vaughan, 1919 P. portoricensis (Vaughan, 1919) P. toulai Vaughan, 1919 — P. baracoaensis P. trinitatis Vaughan, 1926 P. waylandi, nom. nov. AAQAAAAAA а> ле SU CU ed Family PORITIDAE Gray, 1842 The Poritidae are defined as colonial and hermatyp- ic. Colonies are usually formed by extratentacular bud- ding. Corallites (except in the Porites subgenera Syn- araea Verrill, 1864, and Napopora Quelch, 1884) are separated by little or no coenosteum. Thecae are in- distinct and flush with the tissue surface. Septa (except in Alveopora) consist of three to eight loosely-joined, simple, vertical trabeculae forming a meshwork of reg- ularly-arranged pores. Septa are united horizontally by one or more rings of simple synapticulae. The inner- most trabeculae of certain septa are sometimes well- developed forming paliform lobes (sensu Vaughan and Wells, 1943), which have been termed “pali” by most poritid workers. A single columellar trabecula is usu- ally present. In thin-section, the calices appear shallow and obscure, and the internal skeleton consists pri- marily of a lattice-work or reticulum composed of ver- tical rods (trabeculae) and horizontal bars (septal con- nections and synapticulae). DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 75 Genus PORITES Link, 1807 Porites Link, 1807, p. 162. Neoporites Duchassaing and Michelotti, 1866, p. 191. Cosmoporites Duchassaing and Michelotti, 1866, p. 193. not Porites Cuvier, 1798, pp. 678-679, according to Article 23b of the International Code of Zoological Nomenclature (see Veron and Pichon, 1982, p. 141). Type species. — Porites polymorphus Link, 1807, p. 163 (= Madrepora porites Pallas, 1766 [in part], pp. 324—325). Recent, Curacao. As discussed by Vaughan (1901) and Bernard (1905), Pallas (1766) synonymized many different branching corals including some non- Porites in his original description of Madrepora porites. The coral that Vaughan (1901) cited as actually rep- resenting Porites porites is the pre-Linnean Corallium, Doris stellatis of Seba (1756, t. III, р. 202, pl. CIX, fig. 11). The location of Seba’s original specimen is not known. A specimen described as topotypic is figured in Vaughan (1901, pl. 28) and in Vaughan and Wells (1943, pl. 23, fig. 2). Diagnosis. —Colonies massive, ramose, columnar, or encrusting. Corallites usually less than 2 mm in diameter. Septa usually 12 in number, arranged in two cycles following a definite bilaterally-symmetric for- mula consisting of a dorsal septum, a ventral septal triplet, and four lateral septal pairs (as described by Bernard, 1905). Septa composed of one to four tra- beculae, the innermost of which may form pali when thickened. Trabeculae always present in the corallite wall. A single trabecula often forms the columella. Remarks. —The genus Porites occurs worldwide to- day in shallow, clear tropical waters and consists of more than 100 described species. It has been one of the most important reef-building corals since the be- ginning of the Miocene. It is abundant in all reef zones ranging from shallow nearshore areas across the reef crest to the deep forereef (Goreau, 1959), and it dom- inates certain nearshore areas and intermediate fore- reef zones (Geister, 1983). As many as five modern species are commonly recognized throughout the Ca- ribbean region (Goreau and Wells, 1967; Wells and Lang, 1973), with the possibility of one additional Species in Brazil (Verrill, 1901b). The genus is believed to have evolved during the Eocene as a result of the reduction of the third septal cycle in Goniopora (Ber- nard, 1903). Approximately 20 names have been pro- posed for Caribbean representatives during the Neo- gene. Porites baracoaensis Vaughan Plate 16, figures 1-13; Plate 17, figures 1-7; Plate 18, figures 1—4; Text-figures 2—5, 10, 12, 14, 16, 17 Porites baracoüensis Vaughan, 1919, pp. 499-500, pl. 147, figs. 1, la. Porites baracodensis var. matanzasensis Vaughan, 1919, pp. 500- 501, pl. 147, figs. 2—4. ?Porites douvillei Vaughan, 1919, p. 501, pl. 149, figs. 2, 2a, pl. 151, figs. 1, la. Porites toulai Vaughan, 1919, pp. 501-502, pl. 150, figs. 1-4. Description. —Colony branching (branch thickness = 2.9 to 16.2 mm) to form open bushes or thickets. Ca- lices polygonal in shape; small to intermediate in size (0.9 to 1.7 mm in diameter); shallow in depth (0.3 to 0.6 mm); with narrow, regular spacing. Theca may be elevated; composed of one (rarely two) trabeculae; straight; with weak denticles (25.9 denticles per calice). Septa 12 to 13 in number; composed of two trabeculae that form small irregular surface denticles. Septa ver- tically continuous. Dorsal septum relatively reduced in length. Ventral septal triplet fused. Pali usually five in number, prominent. Columella tubercle small (0.06 mm in diameter), may not be developed. Palar syn- apticular ring moderately-developed. Fossa narrow (0.3 to 0.5 mm). In thin-section, vertical trabeculae and horizontal septal junctions are equally thick and well-developed. Dissepiments and synapticulae are rare. A single, prominent trabecula forms the wall without developing any coenosteal reticulum. Three thick, continuous tra- beculae run down the mid-corallite region: these con- sist of two outermost palar trabeculae and a central columellar trabecula. Two poorly-developed trabecu- lae intervene between the wall and pali and form the septa. Holotype.—USNM 325069 (refigured here: Pl. 16, fig. 4; Pl. 17, fig. 1). Measurements of the holotype. —Means of five cor- allites: CD 1.32, CS 1.68, L1 0.428, L2 0.452, L3 0.460, NN 6.4, NS 12.0, NB 0.0, DS 1.0, DW 14.4, PL 5.2, SR 2.0, WR 1.0, ТІ 0.092, T2 0.120, T3 0.116, SD 0.180, P1 0.016, P2 0.304, P3 0.196, C1 0.000, C2 0.000, CW 0.586, SY 0.178, WT 0.232, CE 0.00, PE 0.38, SE 0.25, WE 0.65, CH 25.7, CT 7.4. Type locality. —USGS locality 3476, Marl, Baracoa, Cuba. Upper Pliocene. Lectotype of douvillei (herein selected). — USNM 325106 (refigured here: Pl. 16, fig. 6). Vaughan (1919, p. 501) selected two cotypes for this species, one of which cannot be found (Vaughan, 1919, pl. 151, figs. 1, 1а). The remaining specimen (Vaughan, 1919, pl. 149, figs. 2, 2a) is therefore selected as the lectotype. Type locality of douvillei.—USGS locality 6016, La Boca Formation, Panama. Middle Miocene. Material. — 105 boxes of fragments from 41 localities representing approximately 60 colonies. 32 specimens measured. Remarks. — Vaughan described three branched Neo- gene species, which he assigned to Porites, all with corallite diameters of about 2 mm: (1) Р. baracoaensis, which is distinguished by its mural shelf, thick septal granules, six thick pali, and the lack of a columella tubercle; (2) P. douvillei, which is distinguished by its forking septa and a columella tubercle; (3) P. toulai, which may lack a wall-ridge, forms series of calices, and has irregular pali and a columella tubercle. Study of the Dominican Republic material shows that these characters are all highly variable and that continuous gradational series can be recognized between all of these species as they were originally defined by Vaughan. Therefore, they are synonymized here. The species douvillei is only questionably synonymized, because its septal bifurcations, septal denticles, and irregular branching morphology superficially resemble those of a thin-walled P. portoricensis. The wall, septal gran- ules, and pali appear thicker in the holotype of bara- coaensis than they do in the Dominican Republic ma- terial. However, this may be the result of differential preservation. In an undated, unpublished manuscript, Vaughan (date unknown) described one additional species from the Dominican Republic (P. mauryi), which also be- longs to P. baracoaensis as defined herein. Specimens at the USNM labelled as Р. mauryi in Vaughan’s hand- writing (USNM 63226) appear identical to the NMB Dominican Republic cluster. Re-examination of the Tertiary Puerto Rican corals described by Coryell and Ohlsen (1929) shows that only one specimen belongs to P. baracoaensis: AMNH 23085 (P. toulai) from the Lares Formation. Porites chipolanum Weisbord, 1971 (pp. 20-22, pl. 4, figs. 7, 8, pl. 5, figs. 1, 2) (FSU СН5-5а) resembles P. baracoaensis in branching morphology and cerioid wall structure. However, the larger, deeper calices, five prominent pali, and small columella tubercle of P. chi- polanum are more strongly suggestive of the modern Caribbean species P. porites (Pallas, 1766). No specimens of P. baracoaensis as defined herein appear to be represented in the Frost and Langenheim (1974) material from Chiapas, Mexico. Specimens of P. douvillei (UIX3565-X3571) from the Oligocene La Quinta Formation of Chiapas are distinctly different from Vaughan’s (1919) P. douvillei and from the NMB P. baracoaensis in that they have larger, shallower cor- allites, more septa, more pali, more (although thinner) synapticulae, larger columellae, and thicker walls. Variability.—Porites baracoaensis is probably the least variable of all species studied. Although some- times less than 5 mm, branch thickness averages a constant 8 to 10 mm throughout the section studied (Text-fig. 5a). Although corallite diameter decreases most frequently upsection, it shifts in direction spo- radically yielding no definite trend (Text-fig. 4b). On BULLETIN 325 the other hand, wall structure remains fairly constant upsection (Text-fig. 4a). Within a colony, corallites ap- pear larger and more heavily calcified near the colony base. Comparison. — Porites baracoaensis is distinctly dif- ferent from all the other Neogene Dominican Republic Porites because of its small, shallow corallites, its thin, more solid walls, its prominent pali, and its uniform branching morphology. It somewhat resembles the modern Caribbean species P. furcata Lamarck, 1816, in its branching morphology and wall structure. The major difference between the two species lies in the shallower calices with more prominent columella tu- bercles in P. baracoaensis. Statistical tests, therefore, show that the two species are clearly distinct (Text-fig. 17). As shown on Text-figure 16a, only one Miocene Mediterranean species (P. mancietensis Chevalier, 1961) is similar to P. baracoaensis. Like P. baracoaen- sis, P. mancietensis has fairly small corallites, thin solid walls lacking coenosteal reticulum, shallow calices, and well-developed pali. Porites mancietensis, however, forms small encrusting knobs, whereas P. baracoaensis is branching with uniform long, thin branches. Occurrence. —Rio Cana: Cercado Formation (NMB locs. 16853, 16856, 17004), Gurabo Formation (NMB locs. 16826, 16861, 16863, 16881), and Mao For- mation (NMB locs. 16884, 16885). Rio Gurabo: Gu- rabo Formation (NMB locs. 15806, 15808, 15814, 15837, 15838, 15839, 15840, 15842, 15844, 15845, 15846, 15849, 15850, 15851, 15853, 15856, 15858, 15859, 15862, 15863, 16811, 16882, 16883, 16921, 16934) and Mao Formation (NMB locs. 15822, 15824, 15826, 15830, 15834, 16122). Rio Mao: ?Gurabo For- mation (NMB loc. 16911). Distribution. — Porites baracoaensis has been found widely distributed across the central Caribbean during Neogene time. It occurs outside the Dominican Re- public in the following strata: (1) Lower Miocene Lares Formation of Puerto Rico; (2) Middle Miocene La Boca Formation of Panama; and (3) Upper Pliocene La Cruz and Baracoa formations of Cuba, Bowden Formation of Jamaica. f Porites convivatoris, new species Plate 19, figures 1-13; Plate 20, figures 1—4, Text-figures 2—5, 10, 12, 14, 16, 17 Etymology of the name. – 1. convivator = host; gen- itive singular. Description. —Colony ellipsoidal; small (length = 5.5 to 14.8 mm, width = 3.7 to 9.2 mm); smooth surface; a calcareous tube (approximately 2 mm in diameter) always extends down the long axis of the colony. The tube is 0.2 to 0.5 mm in thickness with faint concentric growth layering and is minutely porous in texture. The DOMINICAN REPUBLIC NEOGENE. 3: FOSTER Pr epitheca ofthe coral extends to the tube edges wherever the tube protrudes beyond the outer colony surface. Calices polygonal, intermediate in size (1.0 to 1.6 mm in diameter), deep (0.6 to 1.0 mm) with narrow, regular- spacing. Theca sinuous, often zigzag in outline, com- posed of one to two trabeculae with prominent den- ticles (mean = 25.3 denticles per calice). Septa 12 in number, composed of two non-palar trabeculae that form small, irregular surface denticles. The uppermost septal surface consists of horizontal flakes, often bi- furcating near the wall. Dorsal septum relatively re- duced in length. Ventral septal triplet fused. Pali weak, irregular, three to five in number. Dorsal palus absent. Columella tubercle present, weakly-developed. Palar synapticular ring poorly-developed. Fossa narrow (0.3 to 0.5 mm). In thin-section, vertical thecal trabeculae are prom- inent; horizontal septal junctions are thin and poorly- developed; dissepiments are abundant; and synaptic- ulae are common. A single prominent trabecula forms the wall, accompanied by occasional coenosteal retic- ulum. Two or three trabeculae run down the mid-cor- allite region: the two outermost form pali that are ir- regular and discontinuous and the third, when developed, forms the columella, which is solid and continuous. Two poorly-developed trabeculae inter- vening between the wall and mid-corallite region form the septa. Holotype. —NMB D5830 (figured here: РІ. 19, figs. ЮЛ Measurements of the holotype.—Means of five cor- allites: CD 1.52, CS 1.51, L1 0.396, L2 0.568, 13 0.608, NN 6.6, NS 12.0, NB 4.8, DS 1.8, DW 24.0, PL 4.6, SR 2.0, WR 1.0, T1 0.080, T2 0.080, T3 0.120, SD 0.136, P1 0.000, P2 0.140, P3 0.120, C1 0.156, C2 0.088, CW 0.460, SY 0.118, WT 0.168, CE 0.27, PE 0.42, SE 0.52, WE 0.69, CH 9.2, CT 10.0. Type locality. —NMB locality 16852, Río Cana, Cer- cado Formation, Dominican Republic. Upper Mio- cene (Saunders, Jung, and Biju-Duval, 1986, text-fig. 15). Paratypes. -NMB D5831-D5841 (figured here: Pl. 19, figs. 2, 3, 6, 8-13; Pl. 20, figs. 1-4). Material.—191 colonies from 15 localities. Ten specimens measured. Remarks.—This species was described by T. W. Vaughan in an unpublished manuscript under the name Porites dominicensis. The manuscript was recently found in a drawer in the Vaughan collection at the USNM and is now archived at the USGS library in Denver. I have not been able to locate the exact ma- terial on which this description was based. Calcareous tubes similar to those characteristic of this species are formed today by a sipunculid worm, Aspidosiphon corallicola Sluiter, 1889, commensal with the coral Heteropsammia cochlea (Spengler, 1781), which inhabits muddy areas of the Great Barrier Reef (see Goreau and Yonge, 1968). As the worm moves, it carries the coral to new feeding areas, often covering a distance of up to one meter in a 24-hour period. Variability.—Although the colony is consistently small and ellipsoidal in shape (Text-fig. 5b), corallite characters do vary widely following few clear trends. Wall structure remains fairly constant stratigraphically upsection (Text-fig. 4a), whereas corallite diameters tend to decrease very slightly (Text-fig. 4b). Similarly, within colonies, corallite diameters decrease from col- ony center to edge; however, such variation may be more strongly related to ontogeny than to environ- ment. Because of the small size of the colony, other within-colony trends are not apparent. Comparison. — Porites convivatoris is unique among the Neogene Caribbean Porites in its small colony size and its unusual ellipsoidal colony shape with a tube extending down the long axis. It also differs in its highly elevated, intermediate-sized calices, which have thin walls, a small columella, and four to five prominent pali. As shown by Text-figure 17, P. convivatoris has calices that resemble those of at least three modern Caribbean species: the finely-branching P. divaricata Lesueur, 1821, the thickly-branching Р. porites (Pallas, 1766), and the encrusting to massive P. branneri Rath- bun, 1887. However, P. convivatoris clearly differs from these three modern forms in its better-developed col- umellar tubercle, weaker pali, and its thicker, bifur- cated septa. As shown in Text-figure 16a, four described Miocene Mediterranean species all closely resemble P. convi- vatoris: P. collegniana Michelin, 1842, P. incrustans Milne-Edwards and Haime, 1851, P. /eptoclada Reuss, 1872, and P. lobatosepta Chevalier, 1961. All have closely-spaced, intermediate-sized calices with high el- evations and well-developed pali and columellae. The septa have numerous denticles and frequently bifur- cate. None of these Mediterranean species, however, form small ellipsoidal colonies with a tube extending down the long axis. Porites collegniana and P. incrus- tans form intermediate to large encrusting colonies, and P. lobatosepta and P. leptoclada are typically branching. Occurrence. —Rio Cana: Cercado Formation (NMB locs. 16836, 16839, 16844, 16852, 16984, 16986) and Gurabo Formation (NMB loc. 16879). Rio Mao: Cer- cado Formation (NMB locs. 16913, 16915, 16916, 16918, 16922, 16926, 16928, 16929). Distribution. —This species is not known from out- side the Upper Miocene and Lower Pliocene of the Dominican Republic. 78 BULLETIN 325 Porites macdonaldi Vaughan Plate 21, figures 1-8; Plate 22, figures 1-6; Plate 23, figures 1—4; Text-figures 2—5, 10, 12, 14, 16, 17, 19 ?Porites anguillensis Vaughan, 1919, pp. 504-505, pl. 150, fig. 5; not pl. 149, figs. 1, la, 1b. Porites (Synaraea) macdonaldi Vaughan, 1919, pp. 506-507, pl. 152, figs. 1-4. Description. —Colonies columnar, massive, or plate- shaped. Plates 3.5 to 15 mm thick, with faint growth bands at 2- to 3-mm intervals. Epitheca having nu- merous linear striations (0.2 to 0.3 mm thick, spaced 1.5 mm apart, and oriented parallel to the direction of growth). Calices circular in shape; intermediate in size (1.1 to 1.6 mm in diameter) and in depth (0.4 to 0.9 mm); with wide, irregular spacing. Frequently intra- tentacular budding forms corallite rows separated by ridges of coenosteum. Theca composed of three to six trabeculae forming a characteristic costal reticulum; sinuous; with weakly-developed denticles (22.1 den- ticles per calice). Septa 12 to 13 in number; composed of three non-palar trabeculae that form small, irregular surface denticles. Septa vertically continuous; of equal length and thickness. Ventral septal triplet fused. Pali weak, zero to five in number. Columella tubercle usu- ally present, moderately-developed. Palar synapticular ring poorly-developed. Fossa of moderate width (0.4 to 0.6 mm). In longitudinal thin-section, horizontal septal junc- tions are thick and predominant, whereas vertical tra- beculae are weak. The dissepiments and synapticulae are well-developed. The corallite wall consists of one to two vertical trabeculae that intergrade with an ex- tensive, irregular coenosteal reticulum. The columella is formed by a tangle of one to two discontinuous ver- tical trabeculae that can be traced down the corallum. The septa are composed of two weak trabeculae. In tangential sections, an irregular arrangement of thick, relatively solid corallite walls incompletely surrounds groups of elliptical speck-like denticles or lobes. The inner circle of these specks is consistently six in number and surrounds a single similarly-shaped columella tu- bercle. е Lectotype (herein selected). — USNM 72376, one of the original cotypes designated and figured by Vaughan (1919, pl. 152, fig. 1). Paralectotypes consist of three additional cotypes (USNM 325046) designated and figured by Vaughan (1919, pl. 152, figs. 2, 3, 3a, 4). The calices of all four type specimens appear to be distorted considerably by subsequent geologic compression of the reefal unit at the type locality in the Emperador Limestone. A fifth possible specimen of this species, which was not designated as a cotype but was figured by Vaughan (1919, pl. 152, figs. 5, 5а), has not been found. The lectotype and one paralec- totype are refigured on Plate 21, figures 1, 4 and on Plate 22, figure 1. Measurements of the lectotype. —Means of five cor- allites: CD 1.33, CS 1.89, L1 0.440, L2 0.460, 13 0.512, NN 7.4, NS 12.4, NB 2.0, DS 2.0, DW 23.2 PL 5.0, SR 3.4, WR 2.8, T1 0.108, T2 0.116, T3 0.124, SD 0.120, P1 0.000, P2 0.120, P3 0.116, C1 0.172, C2 0.132, CW 0.522, SY 0.102, WT 0.444, CE 0.229, PE 0.287, SE 0.325, WE 0.688, CH 11.2, CT 55.9. Type locality. — USGS locality 6016, Empire Quar- ry, Emperador Limestone of the La Boca Formation of Panama. Middle Miocene. Material. — 80 boxes of fragments from 20 localities, representing approximately 80 colonies. 14 specimens measured. Remarks.—This species was originally assigned to the subgenus Synaraea Verrill, 1864 by Vaughan (1919), because of its well-developed coenosteal retic- ulum. However, its calices are distinctly larger (1 to 1.5 mm in diameter) than other modern representa- tives of Synaraea, which usually have calices less than 1 mm in diameter. Additionally, the pronounced coe- nosteal ridges between corallites on the fossil speci- mens are more characteristic of the subgenus Napopora Quelch, 1884, than they are of Synaraea, which tends ртт ры nc... Text-figure 18. —Surface of a modern colony of Porites astreoides having a Synaraea-like appearance, х 16. The corallites of this coral sometimes appear reduced in size and separated by a well-developed wall reticulum in forereef habitats at moderate depths. Wells per- sonal collection #1332, Discovery Bay, 20 m, Jamaica. (Photo cour- tesy of J. W. Wells) DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 79 to develop more superficial corallites that are flush on the surface of the corallum. The subgenus Napopora is poorly known and has never been reported as a fossil or in the Caribbean (Veron and Pichon, 1982). Due to these ambiguities, Р. macdonaldi has not been assigned to a separate subgenus and is herein considered Porites sensu stricto. Results of the canonical variable analyses (Text-fig. 10a) show that P. macdonaldi does not differ enough from P. portoricensis (Vaughan, 1919) or P. waylandi, nom. nov., to warrant separation into a dis- tinct subgenus. Also, some modern Caribbean speci- mens of Porites astreoides Lamarck, 1816, have been found to develop similar small-sized calices separated by extensive coenosteal reticulum in deep, low-light environments (Text-fig. 18). A paratype of P. anguillensis (USNM 325103) is questionably included in the synonymy, because its surface is so badly preserved that corallites can hardly be distinguished. Like the holotype of P. anguillensis (PIU WI-43), it is plate-shaped; however, it does not have the large calices with six prominent pali and the well-developed columella tubercle that are character- istic of the holotype. The paratype appears to have more superficial corallites with a well-developed coe- nosteal reticulum and therefore more likely belongs to P. macdonaldi. The Oligocene Chiapas (Mexico) specimen of Frost and Langenheim (1974) referred to as Porites (Syn- araea) macdonaldi (UCMP 10239, pls. 7-8, figs. 4-5), is lost. From the description and photographs given by Frost and Langenheim (1974), their specimen ap- pears to belong to Р. macdonaldi as described here. Variability. — Although the colony shape is uniform- ly plate-like (Text-fig. 5b), all corallite characters (es- pecially pali number) (Text-fig. 14c) do vary moder- ately, however following no significant trends. Wall thickness appears to decrease slightly upsection strati- graphically (Text-fig. 4a) accompanied by slight in- crease in corallite diameter (Text-fig. 4b). Within col- onies, the coenosteal reticulum is better-developed along the colony edge. Comparison. — Porites macdonaldi is not similar to any other modern Caribbean or Miocene Mediterra- nean species. As explained in the “Comparison” sec- tion for P. portoricensis, it most closely resembles the Neogene Caribbean species P. portoricensis and P. waylandi. It differs from these two species in its rela- tively small corallite diameter, wide columellar syn- apticular ring, well-developed pali, less pronounced ornamentation, and fewer septa. Occurrence. — Río Cana: Gurabo Formation (NMB locs. 16815, 16823, 16826, 16861, 16881). Río Gu- rabo: Gurabo Formation (NMB locs. 15837, 15841, 15846, 15851, 15852, 15855, 15857, 16811, 16882, 16883, 16921, 16933) and Mao Formation (NMB locs. 15824, 15826). Río Mao: ?Gurabo Formation (NMB loc. 16911). Distribution. — Porites macdonaldi ranges in age from late Oligocene to early Pliocene and has been found widely distributed across the central Caribbean. Out- side of the Dominican Republic, it occurs in the following strata: (1) Upper Oligocene La Quinta For- mation of Chiapas, Mexico; (2) Lower Miocene An- guilla Formation of Anguilla; and (3) Middle Miocene La Boca Formation of Panama. Porites portoricensis (Vaughan) Plate 24, figures 1-15; Plate 25, figures 1-6; Plate 26, figures 1—6; Plate 27, figures 1—4; Plate 28, figures 1-4; Text-figures 2—5, 10, 12, 14, 16-19 not Alveopora fenestrata (Lamarck) Dana, 1846, p. 98. Alveopora fenestrata (Dana) Duncan, 1863, p. 437. Goniopora portoricensis Vaughan, 1919, pp. 495-496, pl. 146, figs. 4, 5. Goniopora clevei Vaughan, 1919, pp. 496-497, pl. 145, figs. 1, 3- 6a; ?pl. 145, figs. 2, 2a. Goniopora cascadensis Vaughan, 1919, pp. 497—498, pl. 146, figs. 6-9. ?Porites (Synaraea) howei Vaughan, 1919, pp. 505-506, pl. 151, figs. 2-4. Goniopora ballistensis Weisbord, 1973, p. 32, pl. 11, figs. 1-3; pl. 10, figs. 4, 5 (seen); ?pl. 12, figs. 1, 2 (not seen). Goniopora matsoni Weisbord, 1973, p. 34, pl. 12, fig. 3; pl. 12, figs. 4—6 (seen); pl. 14, figs. 1-3 (not seen). Description. — Colony irregularly branching or co- lumnar (6 to 52 mm thick) with skirted margins that may form plates. Colony surface generally smooth with occasional lumps or protuberances. Calices polygonal to circular, highly variable in size (1.2 to 2.2 mm in diameter) with intermediate depths (0.3 to 1.0 mm) and variable, and in some specimens irregular spacing. Occasional intratentacular budding forming short cor- allite rows and rare ridges in the coenosteum may give the coral a Napopora-like appearance. Theca com- posed of two to five trabeculae commonly form a costal reticulum; sinuous, sometimes zigzag in outline; with prominent granulations (mean = 24.2 granulations per calice). Septa 11 to 13 in number; composed of two to three non-palar trabeculae, the innermost of which commonly develops a large denticle. Septa relatively thick (>0.1 mm). Septal surface consisting of horizon- tal flakes, commonly bifurcating near the wall. Septa of equal length and thickness. Ventral septal triplet free or fused. Pali poorly-developed, if at all. Columella tubercle well-developed. Palar synapticular ring well- developed. Fossa of moderate width (0.4 to 0.7 mm). In longitudinal thin-section, horizontal septal junc- tions predominate. They are uniformly widely-spaced and well-developed and are interrupted only occasion- 80 BULLETIN 325 ally by thinner, irregular vertical trabeculae. Dissepi- ments and synapticulae are rare. Commonly three tra- beculae form the corallite wall; however, this structure cannot be traced far below the surface into the coral- lum. The columella is formed by an irregular tangle of two to three trabeculae. In tangential section, the septal denticles form a characteristic circle of six to eight oval specks, and the septa are composed of two to three thick, irregular trabeculae. The reticulum at the core of the branch axis is especially porous and dominated by vertical structures. Holotype.—USNM 325061 (refigured here: Pl. 24, hp 1; El. 25, Be. 1). Measurements of the holotype. — Means of five cor- allites: CD 1.42, CS 1.88, L1 0.344, L2 0.400, L3 0.432, NN 7.0, NS 12.2, NB 2.4, DS 32, DW 26.6, PL 5.6, SR 3.4, WR 2.8, T1 0.076, T2 0.072, T3 0.060, SD 0.092, P1 0.068, P2 0.128, P3 0.104, C1 0.212, C2 0.100, CW 0.544, SY 0.080, WT 0.416, CE 0.19, PE 0.25, SE 0.27, WE 0.42, CH 74.3, CT 6.0. Type locality. — USGS locality 3191, Lares Forma- tion, four miles west of Lares, Puerto Rico. Lower Miocene. Lectotype of howei (herein selected).— USNM 72890 (refigured here: Pl. 24, fig. 6). Vaughan (1919, pp. 506- 507) designated three cotypes for this species, one of which has been selected as the lectotype (Vaughan, 1919, pl. 151, fig. 4). The remaining two of his original cotypes, USNM 325113 (Vaughan, 1919, pl. 151, figs. 2, 3), are selected as paralectotypes. Type locality of howei.—USGS locality 6016, La Boca Formation, Panama. Middle Miocene. Material. —133 boxes of fragments from 35 locali- ties, representing approximately 60 colonies. 30 spec- imens measured. Remarks. —Study of the Dominican Republic ma- terial shows that this species is highly variable. Con- sequently, three and possibly four of Vaughan's (1919) original Neogene Caribbean species of the Poritidae are synonymized. Vaughan originally described three of these species as belonging to the genus Goniopora Blainville, 1830, because bifurcations of the first two septal cycles suggested the presence of a third poorly- developed septal cycle. However in the Dominican Republic material, these bifurcations appear to be re- lated to decreased calcification in the centers of hori- zontal septal flakes near the wall and not to the actual development of tertiary septa. Similar horizontal flakes have been found to develop in modern Caribbean spec- imens of P. astreoides Lamarck, 1816 in low-light en- vironments (Text-fig. 19). The overall septal arrange- ment also clearly follows a Porites pattern, consisting of four lateral septal pairs and a ventral septal triplet. The Goniopora pattern of fusion of the tertiary septa with the secondary septa cannot be recognized. Bifur- cations can be seen in both primary and secondary septa (see Pl. 28, figs. 1, 2). The four Vaughan species (portoricensis, clevei, cas- cadensis, and howei) were all originally described as branching forms with large corallite diameters (approx. 2 mm), variably-developed mural reticula, thick gran- ules or denticles on the upper surface of septal trabec- ulae, and a dense columella. The species portoricensis was originally characterized by thinner, more delicate septal denticles and columellae. In clevei, the upper surface of the innermost septal trabeculae were de- scribed as thickened to form pali. In cascadensis, three prominent rows of septal granules were developed and the mural reticulum was reduced in thickness, in some specimens forming a zigzag wall ridge. Study of the Dominican Republic material shows that all of these characters vary continuously in P. portoricensis, even across the surface of a single colony, and that they therefore do not warrant species-diagnostic status. Fur- thermore, although it is not complete, the canonical variable polygons of the types of portoricensis, clevei, and cascadensis clearly overlap with the Dominican Republic portoricensis polygon (Text-fig. 10). Unfor- tunately, the holotype of P. portoricensis has been com- pressed due to geologic compaction and lies statisti- " E with flaky septa, x 16. The septa in this coral appear to be formed by horizontal flakes in deep-water habitats. Wells personal collection #2241, Discovery Bay forereef, 42 m, Jamaica. (Photo courtesy of J. W. Wells) DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 81 cally at the margin of the Dominican Republic portoricensis cluster. Therefore, it makes a poor rep- resentative of the species. Vaughan’s species howei is questionably synony- mized with the Dominican Republic portoricensis ma- terial. The type material for howei is poorly preserved and distorted by geologic compaction. The cluster of its types is discrete, but it is nevertheless close enough to the Dominican Republic group still to represent the same species. It differs from the Dominican Republic cluster by having smaller calices (mean corallite di- ameter = 1.134 mm) and a more extensive reticulum. Six pali are commonly developed by the innermost septal trabeculae. The species howei forms subpalmate branches with carinae that may be characteristic of the Dominican Republic specimens (e.g., NMB D5796, Pl. 24, fig. 7; Pl. 26, fig. 3). The statistical analyses show that a paratype of Weis- bord’s (1973) species ballistensis (USNM 68314) clear- ly falls within the cluster of clevei types, and the ho- lotype of matsoni (USNM 68315) falls within the cluster of Dominican Republic portoricensis (Text-fig. 10); therefore, both Weisbord species are included in the synonymy. Similarly, Duncan’s (1863) specimen of A/- veopora fenestrata [BM(NH) R28828] falls within the Dominican Republic portoricensis cluster and is also synonymized. Re-examination of the Tertiary Puerto Rican corals described by Coryell and Ohlsen (1929) shows that specimens of five described species belong to P. por- toricensis: AMNH 23085 (P. toulai Vaughan, 1919), AMNH 23054 (G. portoricensis), AMNH 23042 (G. clevei), and AMNH 23078 (P. douvillei Vaughan, 1919) from the Lares Formation, and AMNH 23076 (P. bar- acoaensis Vaughan, 1919) from the Ponce Formation. Frost and Langenheim’s (1974) figured material of P. baracoaensis (pl. 77, figs. 1-5, UI X3592-X3596) and of Goniopora cascadensis (pl. 79, figs. 3-7, UI Х3598-Х3599) has been lost. However, the photos and descriptions of these two species closely conform to the concept of P. portoricensis described herein. Variability. — Porites portoricensis is the most vari- able of all the species studied; however, consistent trends of variation are extremely complex and there- fore difficult to recognize. Colony morphology, coral- lite size, wall structure, and the development of the coenosteal reticulum all vary widely. Within colonies, corallites are smaller and the coenosteal reticulum bet- ter-developed in thicker columns near the colony base. Only insignificant trends can be detected up the strati- graphic section. Branch thickness increases somewhat upsection (Text-fig. 5) accompanied by a correspond- ing decrease in corallite diameter and slight decrease in wall thickness (Text-fig. 4). Comparison.—This species appears to be unique among branched Neogene Caribbean Porites by having large corallites, bifurcated septa, a thick mural retic- ulum, and thickened septal denticles and/or pali. It appears to be most closely related to the two massive species of Porites represented in the Dominican Re- public material; however, it differs from both P. way- landi and P. macdonaldi by having larger corallites, thicker septa, and numerous prominent septal denti- cles. Porites portoricensis does not resemble any Mio- cene Mediterranean species or any modern Caribbean species. Occurrence. — Rio Cana: Cercado Formation (NMB locs. 16853, 16856), Gurabo Formation (NMB locs. 16815, 16823, 16826, 16881), and Mao Formation (NMB locs. 16884, 16885). Rio Gurabo: Gurabo For- mation (NMB locs. 15808, 15837, 15838, 15840, 15845, 15849, 15850, 15851, 15853, 15855, 15893, 16184, 16811, 16882, 16883, 16921, 16933, 16934) and Mao Formation (NMB locs. 15821, 15822, 15824, 15826, 15834). Rio Mao: ?Gurabo Formation (NMB loc. 16911). Rio Yaque del Norte: Tabera Group (NMB locs. 16909, 17279), Baitoa Formation (NMB loc. 17270), Distribution. —P. portoricensis ranges in age from late Oligocene to early Pliocene and has been found widely distributed throughout the Caribbean. The occurrences in the Dominican Republic are the youngest reported for the species. It has been found outside the Domin- ican Republic in the following strata: (1) Upper Oli- gocene Antigua Formation of Antigua and La Quinta Formation of Chiapas, Mexico; (2) Lower Miocene Anguilla Formation of Anguilla, Santa Ana Formation of Chiapas, Mexico, and Tampa Formation of Florida; and (3) Middle Miocene La Boca Formation of Pan- ama and Ponce Formation of Puerto Rico. Porites waylandi, nom. nov. Plate 29, figures 1-4; Plate 30, figures 1-7; Plate 31, figures 1-4; Text-figures 2-5, 10, 12, 14, 16, 17 not Porites collegniana Michelin, 1842, p. 65, pl. 13, fig. 9. Porites collegniana Michelin. Duncan, 1863, p. 437. not Porites panamensis Verrill, 1866, p. 329. Porites panamensis Vaughan, 1919, pp. 503-504, pl. 148, figs. 1-3a (junior homonym). “Porites floridaprima Bernard” of Weisbord, 1973, pp. 28-29 (in part), unfigured mentioned specimen and ?pl. 8, figs. 1-3 (seen); ?pl. 9, figs. 1-4 and ?pl. 10, figs. 1-3 (not seen). Description. — Colony columnar, massive, or plate- shaped, with distinct growth bands at 4- to 6-mm intervals. Colony surface generally smooth with oc- casional lumps or protuberances. Calices polygonal, intermediate in size (1.2 to 1.8 mm in diameter) with intermediate depths (0.4 to 1.0 mm), having inter- mediate, regular spacing. Some theca elevated; com- 82 BULLETIN 325 posed of two to five trabeculae; sinuous, usually zigzag; with moderate-sized denticles (19.6 denticles per ca- lice). Septa 12 in number; composed of two trabeculae that form small to moderate-sized surface denticles. Septa vertically continuous with bifurcations near the wall. Septa of equal length and thickness. Ventral septal triplet fused. Pali poorly-developed, usually four in number. Columella tubercle well-developed (0.1 to 0.3 mm thick). Palar synapticular ring moderately-devel- oped. Fossa relatively wide (0.5 to 0.8 mm). In longitudinal thin-section, vertical trabeculae and horizontal septal junctions are uniformly widely-spaced and equally thin, forming an open network. Synaptic- ulae, dissepiments, and trabeculae around the colu- mella are especially well-developed. Corallites are sep- arated by an even coenosteal reticulum. The wall is formed by a thick vertical trabecula whose extent can be traced through the corallum. The columella is com- posed of three to four thin vertical trabeculae and the septa of two poorly-developed trabeculae. In tangential section, the septa extend to the columellar synapticular ring. The wall is open and continuous with a regular coenosteal reticulum. Holotype.—USNM 325063 (refigured here: Pl. 29, fig. 1; Pl. 30, fig. 1). Measurements of the holotype. — Means of five cor- allites: CD 1.64, CS 1.85, L1 0.564, L2 0.784, 13 0.616, NN 7.0, NS 11.6, NB 1.6, DS 2.00, DW 17.2, PL 5.2, SR 3.0, WR 1.8, T1 0.140, T2 0.132, T3 0.128, SD 0.152, P1 0.072, P2 0.148, P3 0.160, C1 0.168, C2 0.136, CW 0.710, SY 0.148, WT 0.420, CE 0.15, PE 0.36, SE 0.36, WE 0.73, CH 23.6, CT 89.1. Type locality. — USGS locality 6015, Empire Quar- ry, La Boca Formation, Panama. Middle Miocene. Material. — 39 boxes of fragments from 22 localities, representing approximately 30 colonies. 16 specimens measured. Remarks. — Vaughan (1919) originally described and named this Neogene coral Porites panamensis, appar- ently unaware of Verrill’s (1866) description of the living eastern Pacific species Porites panamensis. The fossil clearly differs from the living coral in corallite size and in the structure of the wall and columella. The fossil has distinctively larger corallites, a better-devel- oped coenosteal reticulum, and a thick columella. Therefore, Vaughan's (1919) fossil species is herein renamed “waylandi” after Vaughan’s middle name. Vaughan (1919, and in Vaughan and Hoffmeister, 1926) described three species of massive Miocene Po- rites having corallite diameters of 1.5 to 2 mm and some (although thin) coenosteal reticula: (1) P. pana- mensis Vaughan, 1919, with small to intermediate- sized corallite diameters and an irregular wall struc- ture, apparently produced by intratentacular budding, (2) P. anguillensis Vaughan, 1919, with large corallite diameters and a regular coenosteal reticulum, and (3) P. trinitatis Vaughan, in Vaughan and Hoffmeister, 1926, with small corallite diameters and a thin, solid wall forming no coenosteal reticulum. In addition, *panamensis" was defined as having six pali, thin syn- apticulae, and a thin columella tubercle; anguillensis as forming flexed laminar plates and having a directive triplet that fuses near the columella, numerous syn- apticulae and a well-developed columella tubercle, and five to six prominent pali; and trinitatis as having thin synapticulae, a thin columella tubercle, six pali, and a directive triplet that fuses at the ventral palus. As shown by Text-figures 10b and 12b respectively, trinitatis and anguillensis are statistically different from “panamen- sis". The three species, therefore, are not synonymized. Although the “holotype” [BM(NH) R2343] repre- sents an unusual branching form, most of Weisbord's (1973) specimens of “P. floridaprima Bernard" clearly fallinto the NMB waylandi cluster (Text-fig. 12b). This includes the specimen measured herein (USNM 66220) as well as other USNM specimens labelled by Vaughan as “Porites willcoxi" from USGS locality 3286 (Ballast Point, Tampa, Florida). Vaughan described P. willcoxi neither in publication nor, to my knowledge, in manu- script. Weisbord's (1973) use of Bernard's (1906, p. 71, pl. 12, fig. 2) description to name the species “‘flor- idaprima” is cited in quotation marks here, because Bernard (1906) did not use Linnean nomenclature in describing the coral and therefore Bernard's (1906) name is invalid. Also for the same reason, the new taxa described in Bernard's (1903, 1905, 1906) three mono- graphs on the Poritidae should be placed on the Official Index of Rejected and Invalid Works in Zoological Nomenclature. Re-examination of the Tertiary Puerto Rican corals described by Coryell and Ohlsen (1929) shows that specimens of three described species belong to P. way- landi: AMNH 23081 (P. panamensis) from Lares, AMNH 23074 (P. astreoides Lamarck, 1816) from Guanica, and AMNH 23077 and AMNH 23079 (P. douvillei Vaughan, 1919) from Lares. Frost and Langenheim's (1974) specimens of P. pan- amensis (UI X3537—X3562) are lost. Nevertheless, their description and photos closely resemble waylandi as described herein. In addition, study of some of Frost and Langenheim's (1974) specimens of P. trinitatis (UI X3576 and X3585) shows that their colony shape and wall structure are similar to P. waylandi. However, the corallites are distinctively small (the mean corallite diameter for UI X3583 is 1.16 mm), and the septa, pali, columellae, and synapticulae are particularly thin. Variability.— Porites waylandi has relatively stable corallite characters, the most variable being wall thick- DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 83 ness and the development of the pali. Corallite di- ameter increases slightly upsection (Text-fig. 4b) and changes in colony height (Text-fig. 5b) appear posi- tively correlated with wall thickness (Text-fig. 4a). Within colonies, corallites appear larger and more widely-spaced from colony top to side. Comparison.—As discussed in the “Comparison” section for P. portoricensis (Vaughan, 1919), ofall Neo- gene Caribbean species of Porites, P. waylandi most closely resembles P. portoricensis and P. macdonaldi Vaughan, 1919. It differs primarily in its thick coe- nosteal reticulum with small calices and generally mound-like shape. The pali are notably poorly-devel- oped, and the septa are thin and heavily ornamented and extend to the columellar synapticular ring. As shown in Text-figure 17, P. waylandi slightly re- sembles the modern Caribbean species P. astreoides Lamarck, 1816. It also resembles modern specimens with widely-spaced corallites of P. branneri Rathbun, 1887, and some specimens of the modern species P. verrilli Rehberg, 1883. However, in P. waylandi, the calices are notably shallower, and the columella tu- bercle better-developed than in the three modern species. The columellar synapticular ring is thick but narrow, and all 12 septa extend to it. Unlike P. bran- neri, calices of P. waylandi have only approximately four weak pali. As shown on Text-figure 16a, P. waylandi is similar morphologically to at least three Miocene Mediterra- nean species of Porites (P. calabricae Chevalier, 1961, Р. pachysepta Chevalier, 1961, and Р. maigensis Kühn, 1925), which all have well-developed wall structures and coenosteal reticula. Of these three, P. maigensis most closely resembles P. waylandi in corallite diam- eter, septal arrangement, thinner synapticulae, and the reduced columellae and pali. Porites maigensis, how- ever, typically forms branches and not mounds or plates as does P. waylandi. Occurrence. —Rio Cana: Cercado Formation (NMB locs. 16853, 16855, 16856), Gurabo Formation (NMB locs. 16817, 16819, 16823, 16881), and Mao For- mation (NMB loc. 16885). Rio Gurabo: Gurabo For- mation (NMB locs. 15846, 15878, 15887, 16811, 16933) and Mao Formation (NMB loc. 15826). Rio Y aque del Norte: Baitoa Formation (NMB locs. 16936, 16943, 17272, 17273, 17275, 17277, 17284, 17289). Distribution. —Porites waylandi is widely distributed across the Caribbean from late Oligocene to early Plio- cene time. The occurrences in the Dominican Republic are the youngest reported for the species. It has been found outside the Dominican Republic in the following strata: (1) Upper Oligocene La Quinta Formation of Chiapas, Mexico; (2) Lower Miocene Tampa Forma- tion of Florida, Lares Formation of Puerto Rico, Río Lajas and Santa Ana formations of Chiapas, Mexico; and (3) Middle Miocene La Boca Formation of Pan- ama, and Ponce Formation of Puerto Rico. Genus GONIOPORA Blainville, 1830 Goniopora Blainville, 1830, pp. 359-360. Type species. — G. pendunculata Quoy and Gaimard in Blainville, 1830. Recent. Port Dorey, New Guinea. Diagnosis. —Colonies massive, ramose, or colum- nar, rarely encrusting. Corallites usually greater than 2.5 mm in diameter, separated by a distinctive, al- though poorly-developed coenosteal reticulum. Septa usually 24 in number, arranged in three cycles follow- ing a definite bilaterally-symmetric formula that con- sists of a dorsal and a ventral septum, two free sec- ondary septa, and four lateral septal pairs (as described by Bernard, 1903). All secondary septa are joined on either side by a tertiary septum. Septa composed of four to eight trabeculae, the innermost of which may form pali when thickened. Columella trabecular, spon- gy, and well-developed. Remarks.— The genus Goniopora occurs today only in the Indo-Pacific region and consists of 32 recogniz- able species (Veron and Pichon, 1982). It is especially abundant along the northern shores of Australia, through Indonesia into the South China Sea, and across the Indian Ocean into the Red Sea. Ecologically, Go- niopora is abundant and known to occur in a broad range of reef habitats extending from the lagoon across the reef crest to the forereef slope. The oldest known fossil representative of the genus is from the lower Cretaceous of Crimea. The genus was common in the Eocene, Oligocene, and Miocene of the Mediterranean and Caribbean, and in the Miocene of the Middle East and Indonesia. It apparently went ex- tinct in the Mediterranean at the end of the Miocene during the Messinian salinity crisis and in the Carib- bean during the early Pliocene (Bernard, 1903). There- fore, the occurrences in the Dominican Republic are among the youngest known for the genus in the Ca- ribbean. Approximately 10 species names have been proposed for the Caribbean region during the Neogene. Goniopora hilli Vaughan, 1919 Plate 32, figures 1-6; Plate 33, figures 1-6; Plate 34, figures 1-4; Plate 35, figures 1, 2; Text-figures 2, 3, 6,8, 11, 13, 14 Goniopora hilli Vaughan, 1919, pp. 488-489, pl. 142, figs. 1, la. Goniopora jacobiana Vaughan, 1919, pp. 492-493, pl. 144, figs. 1- 3a. Goniopora canalis Vaughan, 1919, pp. 494-495, pl. 146, figs. 1-2. Goniopora aucillana Weisbord, 1973, pp. 30-32, pl. 33, fig. 1; pl. 34, fig. 1; pl. 35, fig. 1. Goniopora tampaensis Weisbord, 1973, p. 36, pl. 15, figs. 1-2. 84 BULLETIN 325 Description. —Colonies massive, hemispherical or encrusting; often large (19.6 cm in diameter). Colony surface smooth; may have slight irregularities. Growth bands pronounced, at 2- to 3-mm intervals. Calices polygonal, intermediate in size (2.6 to 3.1 mm), mod- erately deep (0.5 to 1.1 mm) with regular, narrow to intermediate spacing. Theca composed of three to five trabecular rows, 0.5 to 0.9 mm thick, crossed by low costae. Coenosteal reticulum narrow and smooth. Sep- ta 23 to 27 in number, composed of three to five non- palar trabeculae that form prominent surface denticles. Primary and secondary septa of equal length and thick- ness (0.06 to 0.1 mm), extending without fusing to the outer palar synapticular ring. Tertiary septa well-de- veloped, long (0.6 to 0.9 mm), fusing with the second- ary septa. Pali weak, irregular, eight to 12 in number. Columella tangle narrow (0.7 to 1.2 mm in total di- ameter). Synapticulae thin, in six to seven rings. Septa slope gradually from the wall to the center of the col- umella tangle, forming a narrow, V-shaped fossa. In thin-section, vertical trabeculae and horizontal septal junctions are equally thick and well-developed; forming abundant circular pores 0.2 to 0.3 mm in di- ameter, especially across the coenosteum and vertically along septal planes. Dissepiments are prominent, oc- curring at 0.5- to 1.5-mm intervals. Synapticulae are common, occurring at 0.4-mm intervals vertically up a corallite. The columella is formed by two to three twisted trabeculae, the septa by four to five trabeculae that curve gently toward the columella, and the theca by three to five irregular, incomplete trabecular rows. Corallites (especially their columellae) can be traced into the center of the corallum; but because the theca is highly porous and irregular and its texture is similar to that of the septa, the margins between corallites are commonly difficult to discern. Holotype.—USNM 325058 (refigured here: Pl. 32, i Pl. 33. fig BD). Measurements of the holotype. — Means of five cor- allites: CD 3.23, CS 3.26, L1 1.136, L2 1.040, L3 0.656, NN 6.6, NS 22.8, F1 0.0, F2 5.2, DS 3.4, DW 46.8, PL 10.8, SR 4.4, WR 4.2, T1 0.092, T2 0.084, T3 0.096, SD 0.132, P1 0.136, P2 0.136, C1 0.200, C2 0.112, CW 0.886, SY 0.110, WT 0.764, CE 0.230, PE 0.382, SE 0.401, WE 1.491, CH 12.2, CT 52.7. Type locality. — USGS locality 6016, Empire Quar- ry, Emperador Limestone Member of the La Boca For- mation, Panama. Middle Miocene. Lectotype of canalis (herein selected). — USNM 72891 (refigured here: Pl. 33, fig. 3). Vaughan (1919, pp. 494— 495) designated three cotypes for this species, one of which has been selected as the lectotype (Vaughan, 1919, pl. 146, fig. 3). The remaining two cotypes, USNM 325052 (Vaughan, 1919, pl. 146, figs. 1, 2), are selected as paralectotypes. Type locality of canalis.—USGS locality 6016, Em- pire Quarry, Emperador Limestone Member of the La Boca Formation, Panama. Middle Miocene. Material. —20 boxes of fragments from 12 localities, representing approximately 15 colonies. Eight speci- mens measured. Remarks. — Vaughan (1919) described three species of Goniopora with corallite diameters of 3 to 4 mm and a thin coenosteal reticulum: (1) hilli, which forms plates and has three to four teeth per septum, (2) ja- cobiana, which forms hemispherical mounds and col- umns and has five to six teeth per septum, and (3) canalis, which forms branches and has six teeth per septum. All three species have poorly-developed pali and columellae and wall thicknesses of approximately 1 mm. The holotypes of hilli and canalis are both poorly-preserved, as is characteristic of much of the material from the Empire Quarry of the La Boca For- mation in Panama. The holotype of hilli is a fragment (possibly only a portion of the upper growth band of a larger colony), and that of canalis is distorted by compression. Study of the Dominican Republic ma- terial (Text-figs. 11, 13) suggests that all three species belong in the same cluster and should therefore be synonymized. Text-figure 11b shows that a fourth species, G. panamensis Vaughan, 1919, is closely re- lated to the first three; however, G. panamensis has a distinctive coenosteal reticulum that commonly forms ridges, which clearly distinguishes it from related forms. Weisbord (1973) described two new species of Go- niopora from the vicinity of Tampa, Florida that also have corallite diameters of 3 to 4 mm: (1) aucillana from the upper Oligocene Suwannee Limestone, which is distinguished by having a thin laminar wall sur- rounded by synapticulae that give it a perforate ap- pearance, and (2) tampaensis from the lower Miocene Tampa Formation, which is distinguished by its small columella and prominent pali. The holotype and only specimen of aucillana (FSU AU-1a) is deposited in Weisbord’s collection at Florida State University. The wall structure of this specimen closely resembles spec- imens of ?G. decaturensis Vaughan, 1919, a species that is questionably synonymized with G. imperatoris Vaughan, 1919. However, the comparatively large cor- allites and small columella of aucillana are more suggestive of hilli, indicating that the diagnostic lam- inar wall structure that developed in both imperatoris and hilli may be a response to the environment. As shown by canonical discriminant analysis (Text-figs. 11, 13), the holotype and only specimen of tampaensis clearly falls within the hilli Dominican Republic clus- ter. Rhodaraea irregularis Duncan, 1863 (p. 426), from the upper Oligocene of Antigua is a species of Go- niopora with a colony shape and corallite diameters xe DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 85 similar to the Dominican Republic hilli cluster, yet it nevertheless differs in wall and septal structure. The species is based on only one worn, totally-silicified specimen whose septa are thinner and more numerous, walls more distinct, corallites more closely-spaced, and pali better-developed than those of the NMB Domin- ican Republic material. Therefore, it should be con- sidered a separate species until more material from Antigua can be studied. Re-examination of the Tertiary Puerto Rican corals described by Coryell and Ohlsen (1929) shows that specimens of four described species belong to G. hilli: AMNH 23051 (G. jacobiana) and AMNH 23048 (G. hilli) from Guanica, and AMNH 23052 (С. panamen- sis Vaughan, 1919) and AMNH 23039 (G. canalis) from the Lares Limestone. Frost and Langenheim (1974) described six speci- mens of G. hilli from the upper Oligocene La Quinta Formation of Chiapas, Mexico, including two figured specimens. These specimens were originally deposited at the University of Illinois but have been lost (R. L. Langenheim, Jr., written commun., 1984). The pho- tographs and description of this material, however, closely match the Dominican Republic specimens. Variability. — Although morphologic variation with- in this species is not particularly extensive, G. hilli shows the most distinctive directional morphologic trends upsection of all the Poritidae studied. Compar- isons between material from the Rio Yaque del Norte section and from the Rio Gurabo and Rio Cana sec- tions suggest that it increases its corallite diameter and its colony height significantly, possibly in response to increased water depth. Similarly, the amount of coe- nosteal reticulum and the thickness of the columella tangle may increase slightly upsection. Increases in cor- allite diameter, wall thickness, and columella thickness and a decrease in calice elevation can also be traced down the side of a colony. Comparison. —Goniopora hilli overlaps slightly in morphology with G. imperatoris, so that intermediates occur occasionally between the two species clusters. The most important distinguishing characteristics of G. hilli are: (1) its larger corallite diameters; (2) its more elevated calices with reduced columella tangles; (3) its thinner, more irregular wall; and (4) its less distinctive coenosteal reticulum. Of the five Neogene Mediterranean species of Go- niopora described by Chevalier (1961), only one, G. turonensis Chevalier, 1961, is at all similar to G. hilli. However, its corallite diameter is smaller, and it forms only small spherical colonies. Occurrence. —Río Cana: Gurabo Formation (NMB loc. 16818). Río Gurabo: Gurabo Formation (NMB locs. 15808, 15837, 15861, 15884, 16921, 16933). Río Yaque del Norte: Baitoa Formation (NMB locs. 16937, 16943, 16944, 17283, 17284). Distribution. — Goniopora hilli ranges in age from late Oligocene to late Pliocene and has been found widely distributed across the northern and central Caribbean. It occurs outside of the Dominican Republic in the following strata: (1) Upper Oligocene Suwannee Lime- stone of Florida and ?La Quinta Formation of Chiapas, Mexico; (2) Lower Miocene Tampa Formation of Flor- ida, Anguilla Formation of Anguilla, and Lares For- mation of Puerto Rico; (3) Middle Miocene La Boca Formation of Panama, Chipola Formation of Florida, and Ponce Formation of Puerto Rico; and (4) Upper Pliocene La Cruz Marl of Cuba. Goniopora imperatoris Vaughan Plate 35, figures 3-7; Plate 36, figures 1—7; Plate 37, figures 1-4; Text-figures 2, 3, 6, 8, 11, 13, 14 not Madrepora daedalea Forskál, 1775, p. 133. ?Alveopora daedalaea var. regularis Duncan, 1863, p. 426, pl. 14, figs. 4a—. ?Goniopora decaturensis Vaughan, 1919, pp. 490-491, pl. 143, figs. las ?Goniopora decaturensis var. silicensis Vaughan, 1919, р. 491, pl. 143, figs. 2, 2a. ?Goniopora decaturensis var. bainbridgensis Vaughan, 1919, p. 491, pl. 143, figs. 3, 3a. Goniopora imperatoris Vaughan, 1919, pp. 493-494, pl. 142, figs. SE Description. —Colonies massive, columnar, hemi- spherical or plate-shaped; often large (>15 cm in di- ameter), but may form small irregular nodules, 5 cm in diameter. Colony surface smooth. Growth bands, pronounced, at 3- to 4-mm intervals. Calices circular, small (1.8 to 2.8 mm in diameter), shallow (0.3 to 1.1 mm in depth) with regular, intermediate spacing. The- ca composed of three to five trabecular rows, crossed by weak costae. Coenosteal reticulum well-developed, with a smooth upper surface. Septa 20 to 29 in number, composed of three to five non-palar trabeculae that form weak surface denticles. Primary and secondary septa of equal length and thickness (0.06 to 0.10 mm), extending without fusing to the outer palar synapticular ring. Tertiary septa well-developed, of intermediate length (0.4 to 0.6 mm) fusing with the secondary septa. Pali well-developed, nine to 12 in number. Columella tangle relatively thick, 0.7 to 0.9 mm in total diameter. Synapticulae thin, in four to seven rings. Fossa U-shaped, extending one-half to two-thirds of the dis- tance across the corallite. In thin-section, horizontal septal junctions occur more frequently than vertical trabeculae. Trabeculae are thicker than septal junctions, although both are relatively thin. Circular pores (0.1 to 0.2 mm in di- ameter) are abundant, especially across vertical septal planes. Dissepiments are prominent, occurring at 0.7- 86 BULLETIN 325 to 1.2-mm intervals. Synapticulae are common but poorly developed, occurring at 0.3-mm intervals ver- tically up the corallite. The columella is composed of two to three twisted trabeculae, the septa by three to four trabeculae that gently curve toward the columella, and the theca by five or more irregular, incomplete trabecular rows that commonly form a spongy coe- nosteal reticulum. Corallites can be traced to the center of the corallum. The thecal margin itself is relatively distinct and complete. Holotype. — USNM 325049 (refigured here: Pl. 35, fig. 5; Pl. 36, fig. 7). Measurements of the holotype. — Means of five cor- allites: CD 2.26, CS 2.80, L1 0.840, L2 0.660, L3 0.476, NN 6.4, NS 23.2, F1 1.6, F2 5.8, DS 4.0, DW 32.4, PL 8.8, SR 5.2, WR 3.8, T1 0.088, T2 0.072, T3 0.072, SD 0.116, P1 0.096, P2 0.140, C1 0.188, C2 0.080, CW 0.818, SY 0.086, WT 0.760, CE 0.287, PE 0.306, SE 0.344, WE 0.650, CH 54.1, CT 49.2. Type locality. —USGS locality 6016, Empire Quar- ry, Emperador Limestone Member of the La Boca For- mation, Panama. Middle Miocene. Material.—18 boxes of fragments from eight local- ities, representing approximately 13 colonies. Seven specimens measured. Remarks. — Vaughan (1919) described two new species of massive Goniopora from the Caribbean Neo- gene with corallite diameters of 1.5 to 3 mm and well- developed columellae: (1) decaturensis, which forms plates and has a membraniform wall structure and (2) imperatoris, which forms lobate columns and has a well-developed coenosteal reticulum. In addition, two varieties of decaturensis are described: one with larger corallites (2.5 to 4 mm) named var. silicensis, and the other forming small, rounded masses named var. bain- bridgensis. Study of the Dominican Republic material (Text-fig. 11) suggests that the two species are very similar, with the Dominican Republic material more closely resembling imperatoris. Nevertheless, the sig- nificance of the membraniform wall structure in de- caturensis remains unclear without study of additional material from the Chattahoochee Formation of De- catur County, Georgia. Therefore, decaturensis is only questionably synonymized. Because of their larger cor- allites, the two varieties appear almost intermediate between imperatoris and hilli, although their thicker, more delicate intercorallite reticulum is more indica- tive of imperatoris. Duncan (1863) described two new varieties and one new species of Goniopora from the Oligocene of Anti- gua with corallite diameters of 1.25 to 2.5 mm: (1) Alveopora daedalaea var. regularis with larger coral- lites and a coenosteal reticulum; and (2) Alveopora dae- dalaea var. minor Duncan, 1863, and (3) Alveopora microscopica Duncan, 1863, both with smaller calices and a lamellate wall structure. All three types are com- pletely silicified and thus poorly-preserved. Varieties minor and microscopica have almost identical corallite morphologies. They differ from imperatoris in their more solid, straight walls that lack coenosteal reticu- lum, and on further examination ofadditional material these varieties may prove to represent a distinct species. On the other hand, var. regularis appears very close to imperatoris, however, because of the poor preser- vation of the type material, it is only questionably synonymized. In Vaughan’s collections at the USNM, there is a specimen (USNM 63251) from the Baitoa Formation of the Río Yaque del Norte of the Dominican Republic (USGS loc. 8668), which clearly falls in the imperatoris cluster as defined herein. Vaughan identified this coral as Goniopora calhounensis, a name which he never formally described, even in an unpublished manu- script. Later, Weisbord (1971) used the name in de- scribing another species of Goniopora from the Chipola Formation of Florida. Weisbord’s species is described later in this paper. In the middle Miocene Ponce Formation collections described by Coryell and Ohlsen (1929), specimens of four species belong to G. imperatoris as currently de- fined: (1) AMNH 23049, Goniopora imperatoris; (2) AMNH 23044, Goniopora decaturensis, and AMNH 23046, Goniopora decaturensis silicensis; (3) AMNH 23056, Goniopora regularis, (4) AMNH 23048, Go- niopora hilli; and (5) AMNH 23082, Porites pana- mensis Vaughan, 1919. These specimens have gener- ally thinner coenosteal reticula, sometimes forming straight, lamellar wall ridges between calices. These morphologic differences may represent a response to shallower, less turbid marine conditions. Study of Frost and Langenheim’s (1974) specimens of Goniopora imperatoris from the lower Miocene San- ta Ana Formation of Chiapas, Mexico (UI X3606, X3607) reconfirms that they belong to G. imperatoris. Their specimen of Goniopora regularis from the upper Oligocene La Quinta Formation of Chiapas (UI X3538) has been lost; however, the photographs (pl. 80, figs. 4-7) and description (pp. 234-236) closely resemble an imperatoris with reduced coenosteal reticulum. Variability. —Study of the Dominican Republic С. imperatoris shows that variability is reduced in this species. The canonical variable cluster is relatively tight (Text-fig. 11), and low standard deviations were found in single characters (Text-fig. 15, Appendix Ib). Sig- nificant unidirectional trends can only be detected up- section in colony shape, which shows a slight increase DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 87 (Text-fig. 6). Subjective examination of within-colony variability shows that corallites become slightly small- er and the wall thinner from colony top to side. Comparison.—Goniopora imperatoris overlaps slightly with G. hilli as discussed in the Comparison section for G. hilli. However, it resembles no other Neogene Caribbean or Mediterranean species. It is similar morphologically to G. microscopica (Duncan, 1863), from the Oligocene of Antigua, as described above in Remarks. Two Eocene species of Goniopora from the Carib- bean that are close to G. imperatoris include G. taberi Wells, 1934 and G. copoyensis Frost and Langenheim, 1974. Goniopora taberi differs from G. imperatoris by having a ramose growth form and styliform columella. Goniopora copoyensis has a similar well-developed coenosteal reticulum; however, its corallites have no- tably smaller diameters (1.2 to 1.5 mm) and fewer septa than are characteristic of G. imperatoris. Occurrence. —Rio Cana: Cercado Formation (NMB locs. 16853, 16856), Gurabo Formation (NMB loc. 16818), and Mao Formation (NMB loc. 16876). Rio Mao: ?Gurabo Formation (NMB loc. 16911). Rio Ya- que del Norte: Baitoa Formation (NMB locs. 16939, 16944, 17284). Distribution. — Goniopora imperatoris ranges in age from early Miocene (possibly late Oligocene) to early Pliocene and has been found widely distributed across the central and possibly northern Caribbean. The oc- currences in the Dominican Republic are the youngest reported for the species. It has been found outside the Dominican Republic in the following strata: (1) Upper Oligocene ?Antigua Formation of Antigua and ?La Quinta Formation of Chiapas, Mexico; (2) Lower Mio- cene Anguilla Formation of Anguilla and ?Chattahoo- chee Formation of Georgia; and (3) Middle Miocene La Boca Formation of Panama and Ponce Formation of Puerto Rico. Goniopora calhounensis Weisbord Plate 38, figures 1—3; Text-figures 11, 15, 16 Goniopora calhounensis Weisbord, 1971, pp. 18-20, pl. 4, figs. 5, 6. Description. — Colonies massive, hemispherical, large (10 cm in maximum diameter). Colony surface smooth. Calices polygonal, large (3.8 to 5 mm in diameter), shallow (0.65 to 0.75 mm in depth) with regular, wide spacing. Theca composed of four to five trabecular rows, 1 mm thick. Coenosteal reticulum well-devel- oped, with a smooth upper surface. Septa 26 to 27 in number, composed of four to five non-palar trabeculae that form prominent surface denticles. Primary and secondary septa of equal length and thickness (0.14 to 0.2 mm) extending without fusing to the outer palar synapticular ring. Tertiary septa, well-developed, long (1.1 to 1.4 mm) fusing with the secondary septa. Pali well-developed, 12 to 13 in number. Columella tangle relatively thick (1.3 to 1.4 mm in total diameter). Syn- apticulae moderately thick (0.11 to 0.16 mm) in six to eight rings. Septa slope gradually from the wall to the center of the columella tangle, forming a narrow V-shaped fossa. Holotype. —FSU CH3-3a, Weisbord collection, De- partment of Geology, Florida State University, Tal- lahassee, Florida (refigured here: Pl. 38, fig. 1). Measurements of the holotype. — Means of five cor- allites: CD 4.91, CS 6.57, L1 1.42, L2 1.31, L3 1.24, NN 7.2, NS 25.6, F1 0.0, F2 3.6, DS 6.4, DW 50.0, PL 16.4, SR 7.0, WR 6.2, T1 0.13, T2 0.10, T3 0.12, SD 0.24, P1 0.20, P20.21, C10.22, C20.14, CW 2.42, SY 0.14, WT 1.47, CE 0.24, PE 0.24, SE 0.44, WE 0.70, СН 9.1, СТ 27.0. Type locality.—Tenmile Creek, one mile southeast of bridge on state road 73, Calhoun County, Florida. Chipola Formation, Lower Miocene. Material.—Two colonies from two localities. Two specimens measured. Remarks. — Weisbord (1971) originally described this species as consisting of fragments in the Florida State University collection and a few specimens at the USNM and the ANSP. The specimens at the USNM bearing Vaughan's label **Goniopora calhounensis" clearly be- long to G. imperatoris as discussed earlier in the Re- marks section for that species. The ANSP specimen (ANSP 10933), which Weisbord described as typical of G. calhounensis, belongs instead to Goniopora hilli. I have been unable to locate any other material. This leaves only three specimens (the holotype and the two NMB Dominican Republic specimens) that actually belong to the species. Variability. — Because of the poor preservation and scarcity of material, it is difficult to assess variability within this species. Wide variability has been noted in corallite diameter, septum length, and total number of septa (Text-fig. 15); however, subjective examination of these characters shows no distinctive trends that can be related to environment. Therefore, variation in size- related characters may be caused by ontogeny. Little variation is noted in wall thickness or the coenosteal reticulum (Text-fig. 15). Comparison. — Goniopora calhounensis is markedly distinct from all other Tertiary Caribbean Goniopora including G. hilli Vaughan, 1919, and G. imperatoris Vaughan, 1919, because of its large, flat corallites with well-developed columella tangles. As shown in Text- figure 11, no overlap is found with the first two Go- niopora species. Two species described by Chevalier (1961) from the Burdigalian of the Aquitaine Basin (St. Paul-les-Dax, France), however, closely resemble G. calhounensis in these characters: (1) “Goniopora daxi- tertia Bernard” and (2) Goniopora globulosa Chevalier, 1961. The only distinctive difference between calhou- nensis and these two Mediterranean species is in wall structure which appears to be wider, less porous, and better-developed in G. calhounensis. Occurrence. —Rio Cana: Mao Formation (NMB loc. 16875). Rio Gurabo: Gurabo Formation (NMB loc. 16921). Distribution.—G. calhounensis appears to be re- stricted to the Miocene and lower Pliocene of the northeast Caribbean. The only occurrence outside the Dominican Republic is in the lower Miocene Chipola Formation of Florida. Genus ALVEOPORA Blainville, 1830 Alveopora Blainville, 1830, pp. 358-359. Type species. — Madrepora daedalea Forskäl, 1775. Recent. Red Sea. The holotype of this species has been lost and there is some doubt as to whether it actually belongs to the genus Alveopora as now conceived by many coral workers (see Veron and Pichon, 1982). Diagnosis. — Colonies massive or ramose. Corallites circular or polygonal, mostly juxtaposed. Septa usually 12 in number, having no regular pattern of fusion, and composed of spines that project inward from the theca. Vertical septal trabeculae rarely developed. Theca composed of 12 trabeculae linked by horizontal syn- apticulae to form a lattice-like arrangement of pores. Remarks. —The genus Alveopora occurs today only in the Indo-Pacific region and consists of approxi- mately 15 described species (Veron and Pichon, 1982). It has been recorded from as far east as Hawaii and Samoa across Micronesia and Indonesia through the Maldives to the Red Sea. Veron and Pichon (1982) have recognized seven species in eastern Australia. Ecologically, Alveopora occurs in a range of reef hab- BULLETIN 325 itats; however, it is seldom abundant, and species dis- tributions are usually narrowly confined. The oldest known fossil representative of the genus is from the Eocene of Madagascar. The genus is well- known in Europe during the Oligocene and Miocene but it occurs only rarely in the Caribbean during the Oligocene, Miocene, and Pliocene. Only two names, Alveopora chiapanecae Frost and Langenheim, 1974 and Alveopora tampae Weisbord, 1973, have been pro- posed for the Caribbean region during the Tertiary. Alveopora tampae Weisbord Plate 38, figures 4—7 Alveopora tampae Weisbord, 1973, pp. 37-38, pl. 6, figs. 4-6, pl. 7, figs. 4—5. Description. —Colonies massive with even surfaces, usually forming small regular spheres less than 4 cm in diameter. Calices circular, 2 to 3 mm in diameter. Septal spines arranged in two complete cycles with an occasional incomplete third cycle. Theca thick (0.5 mm), composed of one to three trabeculae. Endothecal dissepiments 0.2 mm thick, spaced at 0.8- to 1.2-mm intervals. Thecal pores equal, regularly arranged, cir- cular (some elongate), approximately 0.5 mm in di- ameter, separated by synapticulae approximately 0.2 to 0.3 mm thick. Holotype.—The holotype cannot be found at the USNM. Its catalog number was incorrectly cited by Weisbord (1973) as USGS 2115, which is a locality number. Two topotypes (USNM 66224, 66225) were therefore studied and are figured here on Plate 38, figures 4—6. Measurements of a topotype (USNM 66224).— Means of five corallites: CD 2.42, CS 3.18, L1 0.96, L2 0.29, NN 6.4, NS 12.6, WR 1.5, T1 0.20, T2 0.13, SY 0.19, WT 0.625, CH 28.2, CT 31.9. Type locality. — USGS 2115, 2.5 miles west of Tampa, Florida along the shores of Hillsborough Bay, Tampa Formation. Lower Miocene. Material. — One colony from one locality, deposited at the USNM. DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 89 | Appendix Ia.— Means (+1 standard deviation) of all corallite characters in the five species of Porites herein described. Abbreviations for characters are explained in Table 3. “п” = number of colonies measured. Five corallites were measured in each colony. | Р. convivatoris P. baracoaensis P. macdonaldi P. waylandi P. portoricensis | (n = 10) (n = 32) (n = 15) (n = 15) (n = 30) | CD* I7 (GEO) ED) 12 EO) 1.45 (+0.23) 1.42 (+0.15) 1.69 (+0.28) | CS 1.33 (+0.14) 1.32 (0.17) 1.81 (+£0.26) 1.84 (+0.26) 1.98 (+0.24) | L1* 0.410 (+0.059) 0.398 (+0.082) 0.404 (+0.088) 0.440 (+0.080) 0.554 (+0.094) | 12 0.506 (+0.068) 0.489 (+0.070) 0.493 (+0.083) 0.498 (+0.077) 0.607 (+0.122) | L3 0.521 (+0.080) 0.494 (+0.077) 0.474 (+0.065) 0.523 (+0.091) 0.612 (+0.109) | NN 6.5 (+0.4) 6.5 (0.4) 7.0 (+0.4) 7.4 (0.6) 6:6 (0.3) | NS DA el) 1) 1202202) 11.8 (0.4) 12.0 (+0.4) 12:05 (402) | NB ЭОК (Ge 1:2) 4222 E 1.9 (0.6) 2102 2095) 23:20) | DSt 2.0 (+0.4) 2.3 (+0.4) 23 (04) 2:0 (EOS) 2.8.05) | DW 25:3 tx (ete) 2,9228) 95 CID 220 (+25) 24.1 (+1.5) | PL* 413,95 (220.5) SO ERO) 4.0 (=1.1) ВО (PH 33. (50%) SRT 202602) 2.8 (+0.4) 3.0 (0.4) 3.4 (+0.4) 3.4 (+0.4) | WR* То (ЖОО) 1273202) sale EO) 3:97:07) 30 20:7) | ТІ 0.090 (--0.015) 0.098 (--0.012) 0.106 (--0.020) 0.092 (+0.015) 0.116 (+0.025) | T2 0.091 (+0.021) 0.106 (+0.014) 0.112 (+0.020) 0.094 (0.010) 0.126 (+0.030) | тз 0.098 (+0.021) 0.106 (+0.012) 0.106 (40.016) 0.096 (+0.012) 0.122 (40.025) | SD} 0.105 (+0.014) 0.125 (+0.021) 0.130 (+0.056) 0.116 (+0.015) 0.130 (+0.023) | Р1 0.002 (+0.005) 0.014 (+0.020) 0.021 (+0.020) 0.019 (+0.021) 0.013 (+0.023) | P2 0.087 (+0.023) 0.114 (+0.027) 0.089 (+0.053) 0.050 (+0.037) 0.053 (+0.043) | P3 0.093 (+0.022) 0.111 (+0.016) 0.111 (+0.027) 0.093 (+0.019) 0.101 (+0.029) | С1 0.122 (+0.059) 0.071 (+0.037) 0.177 (0.052) 0.144 (+0.049) 0.138 (+0.040) C2 0.079 (+0.035) 0.059 (+0.031) 0.199 (+0.033) 0.097 (+0.034) 0.103 (+0.030) CW* 0.416 (+0.055) 0.427 (50.053) 0.599 (+0.067) 0.507 (+0.040) 0.555 (+0.080) syt 0.085 (+0.018) 0.113 (50.020) 0.114 (+0.015) 0.094 (+0.012) 0.124 (+0.023) WT* 0.170 (+0.075) 0.120 (+0.030) 0.602 (+0.124) 0.652 (+0.156) 0.508 (+0.158) CE 0.22 (+0.06) 0.14 (+0.08) 0.26 (+0.08) 0.22 (+0.10) 0.21 (+0.06) PE 0.46 (+0.11) 0.37 (+0.41) 0.32 (+0.09) 0.33 (+0.08) 0.31 (+0.08) SE 0.53 (+0.08) 0.35 (+0.07) 0.36 (+0.10) 0.37 (+0.06) 0.43 (+0.10) WE* 0.79 (+0.12) 0.43 (+0.08) 0.65 (+0.12) 0.71 (+0.15) 0.69 (+0.17) * characters whose means and standard deviations аге diagrammed in Text-figure 14. T characters not used in canonical discriminant analyses. Appendix Ib.— Means (+1 standard deviation) of all corallite characters in the three species of Goniopora herein described. Abbreviations for characters are explained in Table 4. “п” = number of colonies measured. Five corallites were measured in each colony. G. imperatoris G. hilli G. calhounensis G. imperatoris G. hilli G. calhounensis (n = 10) (n = 6) (n = 2) (n = 10) (n = 6) (п = 2) CD* 2.19 (+0.30) 2.92 (+0.19) 4.44 (+0.79) T2t 0.076 (20.011) 0.082 (+0.015) 0.174 (0.042) cs 2.49 (+0.22) 3.04 (+0.30) 4.37 (+0.66) T3t 0.064 (+0.010) 0.073 (+0.015) 0.160 (+0.023) L1* 0.734 (+0.106) 1.005 (+0.085) 1.646 (+0.320) SDt 0.109 (+0.013) 0.139 (+0.028) 0.248 (+0.028) L2 0.624 (+0.076) 0.915 (+0.056) 1.370 (+0.167) Р1 0.090 (20.017) 0.112 (40.014) 0.154 (+0.048) L3 0.497 (+0.064) 0.771 (+0.094) 1227220215) P2 0.139 (+0.040) 0.137 (+0.037) 0.208 (+0.040) NN О (BOS) 057 (E02) 6.8 (0.3) C1 0.167 (+0.047) 0.186 (+0.058) 0.212 (+0.006) NS* 2372 (БИІ) 24.3 (1.5) 26.4 (+1.1) C2 0.088 (+0.023) 0.125 (+0.028) 0.160 (+0.034) Fir 0.8 (1.9) 0.6 (+0.4) 0.0 (0.0) CW* 0.871 (+0.095) 0.929 (30.166) 1.350 (+0.062) F2t S.6 (E08) 0 EO) 5.4 (+0.3) SY* 0.077 (+0.007) 0.084 (+0.014) 0.139 (+0.024) DSt 4.3 (0.6) ST GEO) 2487 (£06 WT* 0.647 (+0.093) 0.688 (+0.153) 1.050 (+0.008) DW 33.0 (+3.5) 38.4 (+3.5) 45.3 (+5.8) CE 0.23 (+0.07) 0.26 (+0.05) 0.27 (+0.03) PL* 10.7 (0.9) 9 (ELS 122 (£06) PE 0.31 (+0.10) 0.32 (+0.07) 0.26 (+0.01) SR DEO) 6.9 (0.6) 2 (ЕЗІП!) SE* 0.25 (+0.10) 0.40 (+0.12) 0.29 (+0.05) WR 3.8 (50.4) 3.8 (0.6) 4.5 (+0.4) WE 0.66 (+0.23) 0.76 (+0.22) 0.71 (+0.05) Tif 0.077 (+0.016) 0.091 (+0.020) 0.172 (+0.040) * characters whose means and standard deviations are diagrammed in Text-figure 15. T characters not used in canonical discriminant analyses. 90 BULLETIN 325 Appendix Па. — Canonical discriminant analysis of Porites groups Appendix IIb.— Canonical discriminant analysis of Porites groups 2 and 4. Total-sample correlations between the canonical variable 1, 3, and 5. Total-sample correlations between the canonical vari- and the original variables (COR), and standardized canonical coef- ables and the original variables (COR), and standardized canonical ficients (SCC). Only values with high magnitudes are given. Abbre- coefficients (SCC). Only values with high magnitudes are given. The viations for characters are explained in Table 3. canonical variables are labelled CV 1—2. Abbreviations for characters are explained in Table 3. COR SES NS — (198) Original 29,6; 3 PL 0.534 0.80 variable COR SEE COR SCC WR —0.546 1155 CD 0.458 0.80 54 = T2 — 0.96 cs $ —1.03 $ ‘a х = ines Li —0.656* —2.08* ES s di. : L2 —0.490 0.93 — — 54 M sas L3 —0.592 —0.79 = 0.45 na ee ues NN 0.427 T 0.614* 0.72 А ү WR — О - 0.48 ТІ - - —0.452 — Appendix IIc.— Canonical discriminant analysis of Goniopora T2 em = soos = groups | and 2. Total-sample correlations between the canonical T3 = = —0.438 = variable and the original variables (COR), and standardized canon- CW = 1.28 E3015 16 ul de ical coefficients (SCC). Only values with high magnitudes are given. WT E 0.94 = = Abbreviations for characters are explained in Table 4. PE = = = 0.59 SE - - -— —0.99 Original WE — — = 0.83 variable COR SCC CD 0.802* 1.04* CS 0.640 0.68 L1 0.705 — L2 0.793* E L3 0.800* 0.49 NN — WS DW 0517 -- SR 0.596 — CE - 0.56 Appendix IId.—Canonical discriminant analysis of five fossil and six modern Caribbean species of Porites. Total sample correlations between the canonical variables and the original variables (COR), and standardized canonical coefficients (SCC). Only values with high magnitudes are given. The canonical variables are labelled CV1-5. Abbreviations for characters are explained in Table 3. Original Evil EVA СУЗ СУ4 CV5 variable COR SCC COR SCC COR SCC COR SCC COR SCC CD - O72 - - - - - - - - cs 0.702 — — 0.68 — = = 955 — — L1 - — 0.502* 0.60 — —0.94 — — — — L2 — - = — - - = 061 0.485* - 13 — — — — —0.360 _ = = = = NN — — - - - — 0.484 0.65 — — NB —0.586 — - - - — — 0.45 — - DW 220.916 = — 0.76 = 0,370 = = = = == PL 00527 - - - — — — _ — WR USS 1.14* — 0.69 - bes} = == = 1.10 12 — = 0.456 — — — - 0.64 0.424 - T3 — — 0.463 — -- - - - 0.487% - P2 -0.336 - - — - - - — 0.410 — ES -- - - - - - - - 0.481% = C1 0/595 - - - — - - -0.63 - - CW 0.617 = — = = 0.83 — —0.47 = — WT 0.866* 0.51 — - - 1.25 — 0.69 = жї СЕ — - — — — 0.83 —0.534* = = = SE — — 019195 038 —0.404* - - -0.84% E — WE 0.614 1.02 —0.480 el 202975 IIS — 0.67 — - * most important variables. DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 91 REFERENCES CITED Айту, С. C., and Carrión-Torres, С. 1963. Shallow-water stony corals of Puerto Rico. Caribbean Journal of Science, vol. 3, pp. 133-162, pls. 1-21. Bernard, H. M. 1900. 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Editions de l'Académie bulgare des Sci- ences, Sofia, 472 pp., 161 pls. 94 BULLETIN 325 EXPLANATION OF PLATE 15 Calices of six modern described species of Porites from the Caribbean/Western Atlantic region me 2m... All photos, X 10. The three massive species (figs. 1-3) have traditionally been distinguished on the basis of calice characters such as wall thickness, calice elevation, development of pali, and development of the columella tubercle. The three branched species (figs. 4-6) have been distinguished on the basis of branch thickness: P. porites with swollen branch tips, greater than 20 mm in diameter; P. furcata with dichotomous branching, about 10 mm in diameter; P. divaricata with thin branches, less than 6 mm in diameter. Vaughan (1901) believed only two true species existed in the Caribbean: one represented by figures 1 and 2, the other by figures 3 through 6; whereas, more recent studies such as Brakel (1976, 1977) and Wells and Lang (1973) suggest that there may be as many as four to six. Calices of the Neogene Dominican Republic Porites clearly differ from these modern specimens by having generally larger, shallower corallites. Calices of P. porites (Pallas) in figure 4 and P. divaricata Lesueur in figure 6 slightly resemble one of the fossil species, P. convivatoris (Text- fig. 15). Figure 1. Porites astreoides Lamarck, 1816. Holotype. MHNP, Lamarck Collection No. 169, Mers d'Amérique; distinguished by deep calices, the lack of pali, and a small columella tubercle. 2. Porites verrilli Rehberg, 1893. Holotype. YPM 4539, Abrolhos Reefs, Brazil; like P. astreoides but having extensive coenosteal reticulum, deeper calices, and a prominent columella tubercle. 3. Porites branneri Rathbun, 1887. Holotype. USNM 10961, Parahyba do Norte, Brazil; distinguished by a prominent circle of five pali and no columella tubercle. 4. Porites porites (Pallas, 1766). Figured Specimen. MHNP, Lamarck Collection No. 150, Antilles (holotype of Porites clavaria Lamarck); distinguished by shallow, large calices with six pali and a columella tubercle. 5. Porites furcata var. 2 of Lamarck, 1816. Figured Specimen. MHNP, Lamarck Collection No. 155, Mers d'Amérique; like P. branneri but having smaller calices. 6. Porites divaricata Lesueur, 1821. Figured Specimen. USNM 72377, East of Pulaski Buoy, Florida; like P. porites but having shallower calices. PLATE 15 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 16 DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 95 EXPLANATION OF PLATE 16 Poritesibaracoaensis Naushan . 5. е сас 2 о 7.5 Branch fragments and thin-sections. The colony morphology is fairly uniform, forming bushes with branches | commonly 0.5 to 1.5 cm in thickness. In thin-sections cut perpendicular to branch axes, an irregular core is | seen surrounded by a uniform network of horizontal septal junctions and vertical trabeculae. The vertical | pali, columellae, and mural trabeculae predominate in both transverse and longitudinal thin-sections. Figure 1. Figured Specimen. NMB D5811. Lower Pliocene, NMB locality 16826, Rio Cana, Gurabo Formation, Dominican Republic. Colony fragments as preserved within the sediment matrix, x1. | 2. Figured Specimen. NMB D5812. | Upper Miocene, NMB locality 16883, Rio Gurabo, Gurabo Formation, Dominican Republic. Branch surface, x1. | 3. Figured Specimen. USNM 325105a. | Middle Miocene, USGS locality 6106, La Boca Formation, Panama. Branch surface, x 1 (holotype of Porites toulai Vaughan). 4. Holotype. USNM 325069. Upper Pliocene, USGS locality 3476, Baracoa, Cuba. Branch surface, x1. 5. Figured Specimen. USNM 325067a. Upper Pliocene, USGS locality 3461, Matanzas, Cuba. Branch surface, x 1 (holotype of Porites baracoaensis matanzasensis Vaughan). 6. Possible synonym, lectotype of Porites douvillei Vaughan. USNM 325106. Middle Miocene, USGS locality 6016, La Boca Formation, Panama. Branch surface, x1. 7. Figured Specimen. NMB D5813. Upper Miocene, NMB locality 15850, Rio Gurabo, Gurabo Formation, Dominican Republic. Branch surface, x1. 8. Figured Specimen. NMB D5814. Upper Miocene, NMB locality 16811, Rio Gurabo, Gurabo Formation, Dominican Republic. Branch surface, x1. 9. Figured Specimen. NMB D5815. Upper Miocene, NMB locality 15859, Rio Gurabo, Gurabo Formation, Dominican Republic. Branch surface, x1. 10. Figured Specimen. NMB D5816. Upper Miocene, NMB locality 15849, Rio Gurabo, Gurabo Formation, Dominican Republic. Thin-section cut perpendicular to branch axis, x5. 11. Figured Specimen. NMB D5817. Upper Miocene, NMB locality 16811, Rio Gurabo, Gurabo Formation, Dominican Republic. Thin-section cut perpendicular to branch axis, х5. 12. Figured Specimen. NMB D5817. Same specimen as figure 11 above. Transverse thin-section, X25, showing one complete corallite (upper left) and portions of three others. 13. Figured Specimen. NMB D5818. Upper Miocene, NMB locality 16934, Rio Gurabo, Gurabo Formation, Dominican Republic. Longitudinal thin-section, x 25, through one complete corallite. 96 BULLETIN 325 EXPLANATION OF PLATE 17 Porites:baracoaensis N augha soos т a ЫЕ Сп cio: Close-ups of calical surfaces. Calyx characters vary considerably. A thin, denser wall, low calice elevation, prominent pali, and a small corallite diameter are characteristic. Figure 1. Holotype. USNM 325069. Same specimen as Plate 16, figure 4. Calical surface, x10. 2. Figured Specimen. USNM 325067a. Same specimen as Plate 16, figure 5. Calical surface, x10. 3. Figured Specimen. NMB D5812. Same specimen as Plate 16, figure 2. Calical surface, x 10. 4. Figured Specimen. NMB D5819. Upper Miocene, NMB locality 15849, Río Gurabo, Gurabo Formation, Dominican Republic. Calical surface, x10. 5. Figured Specimen. NMB D5820. Upper Miocene, NMB locality 15856, Río Gurabo, Gurabo Formation, Dominican Republic. Calical surface, x10. 6. Figured Specimen. NMB D5815. Same specimen as Plate 16, figure 9. Calical surface, x10. 7. Figured Specimen. NMB D5813. Same specimen as Plate 16, figure 7. Calical surface, x10. BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 17 ES ا‎ AA € “Ж ‘iar, > LXVI * > wp қ ta іе ae E Жа ж” BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 18 Porites’baracoaensis- Vaughan... в. уиул УИ е DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 97 EXPLANATION OF PLATE 18 SEM photographs of calices. Of all the Neogene Dominican Republic Porites, this species has the most well- developed pali and columella tubercle. The wall commonly consists of a single prominent trabecular row as seen in figures 1 and 2. Figure Il 2 9, Figured Specimen. ММВ 05821. Upper Miocene, ММВ locality 16811, Rio Gurabo, Gurabo Formation, Dominican Republic, х 28. Figured Specimen. NMB D5821. Same specimen as figure 1 above, x67. Figured Specimen. NMB D5822. Upper Miocene, NMB locality 16811, Río Gurabo, Gurabo Formation, Dominican Republic, x 28. . Figured Specimen. NMB D5823. Upper Miocene, NMB locality 15853, Río Gurabo, Gurabo Formation, Dominican Republic, x 28. 98 BULLETIN 325 the long axis. Calices are highly elevated and separated by a single prominent row of mural trabeculae. In thin-section, vertical trabeculae and dissepiments dominate the morphology. Horizontal septal junctions are rare. pa ы EXPLANATION ОЕ PLATE 19 Page FO O OEM HOLD DEW ресе 2-65 о 76 | Colony surfaces and thin-sections. Colonies consist of small ellipsoids. A calcareous tube always extends down > | | Figure 1. Holotype. NMB D5830. Upper Miocene, NMB locality 16852, Rio Cana, Cercado Formation, Dominican Republic. Whole colony, х2. 2. Paratype. NMB D5833. Upper Miocene, NMB locality 16836, Rio Cana, Cercado Formation, Dominican Republic. Whole colony, x2. 3. Paratype. NMB D5832. Lower Pliocene, NMB locality 16879, Rio Cana, Gurabo Formation, Dominican Republic. Whole colony, х2. 4. Figured Specimen. USNM 64669. Upper Miocene, USGS locality 8525, Rio Mao, ?Cercado Formation, Dominican Republic. Whole colony, x2. 5. Figured Specimen. USNM 64669. Same locality as figure 4 above. Whole colony, x 2. . Paratype. NMB D5834. Upper Miocene, NMB locality 16836, Río Cana, Cercado Formation, Dominican Republic. Whole colony, x 2. . Holotype. NMB D5830. Same specimen as figure 1 above. Calical surface, х 10. . Paratype. NMB D5831. Upper Miocene, NMB locality 16928, Rio Mao, Cercado Formation, Dominican Republic. Whole colony, x10. b . Paratype. NMB D5835. | Upper Miocene, NMB locality 16922, Rio Mao, Cercado Formation, Dominican Republic. Thin-section cut perpendicular to the tube, x5. 10. Paratype. NMB D5836. Upper Miocene, NMB locality 16928, Rio Mao, Cercado Formation, Dominican Republic. Thin-section cut perpendicular to the tube, x5. 11. Paratype. NMB D5833. Same specimen as figure 2 above. Calical surface, x10. 12. Paratype. NMB D5837. Upper Miocene, NMB locality 16928, Rio Mao, Cercado Formation, Dominican Republic. Transverse thin-section, x 25, showing one complete corallite. 13. Paratype. NMB D5838. Upper Miocene, NMB locality 16839, Rio Cana, Cercado Formation, Dominican Republic. Longitudinal thin-section, x 25, showing the wall between two corallites. оо - ON © Í ( 1 ! BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 19 “жалт € ж ж BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 20 DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 99 EXPLANATION OF PLATE 20 Porites;convivatorisznew;species m nee 76 SEM photographs of calices. Calices are elevated and have approximately five thin pali and a thin columella tubercle. Septa are usually heavily ornamented. Figure 1. Paratype. NMB D5839. Upper Miocene, NMB locality 16928, Rio Mao, Cercado Formation, Dominican Republic, x28. 2. Paratype. NMB D5839. Same specimen as figure 1 above, x67. 3. Paratype. NMB D5840. Upper Miocene, NMB locality 16836, Río Cana, Cercado Formation, Dominican Republic, x28. 4. Paratype. NMB D5841. Upper Miocene, NMB locality 16928, Río Mao, Cercado Formation, Dominican Republic, x28. BULLETIN 325 EXPLANATION OF PLATE 21 ROLES HACAONGIATN AOI AN see ое 78 Colony fragments and X-radiographs of colonies. The colony morphology consists primarily of plates with growth bands at 1 to 2 mm intervals. Ridges of coenosteal reticulum commonly extend irregularly across the plate surface. Valleys between ridges are formed by series of calices produced by intratentacular budding. X-radiographs show that horizontal septal junctions dominate the internal structure. Figure 1. Lectotype. USNM 72376. Middle Miocene, USGS locality 6016, La Boca Formation, Panama. Fragment of a plate-shaped colony, x1. 2. Figured Specimen. NMB D5842. Upper Miocene, NMB locality 16921, Rio Gurabo, Gurabo Formation, Dominican Republic. Plate-shaped colony, х1. 3. Figured Specimen. NMB D5843. Upper Miocene, NMB locality 15841, Rio Gurabo, Gurabo Formation, Dominican Republic. Plate-shaped colony fragment, x1. 4. Paralectotype. USNM 325046a. Middle Miocene, USGS locality 6016, La Boca Formation, Panama. Colony fragment, х1. 5. Figured Specimen. NMB D5844. Upper Miocene, NMB locality 16933, Rio Gurabo, Gurabo Formation, Dominican Republic. X-radiograph, х1. 6. Figured Specimen. NMB 105845. Upper Miocene, NMB locality 16911, Río Mao, ?Gurabo Formation, Dominican Republic. Plate-shaped colony, x1. 7. Figured Specimen. NMB D5846. Upper Miocene, NMB locality 16883, Rio Gurabo, Gurabo Formation, Dominican Republic. Plate-shaped colony fragment, х1. 8. Figured Specimen. NMB D5847. Lower Pliocene, NMB locality 15826, Rio Gurabo, Mao Formation, Dominican Republic. X-radiograph, х1. BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 21 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 22 DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 101 EXPLANATION OF PLATE 22 Page Porites: macdonaldi Мапа е. 78 Close-ups of calical surfaces and thin-sections. The calical surface is composed of extensive coenosteal reticulum that may form irregular ridges between corallite rows. The calices themselves appear small and superficial. Thin-sections show well-developed horizontal septal junctions, weakly-developed vertical trabeculae, and prominent granulations. Figure 1 2 5). . Lectotype. USNM 72376. Same specimen as Plate 21, figure 1. Calical surface, x5. Figured Specimen. NMB D5846. Same specimen as Plate 21, figure 7. Calical surface, х5. Figured Specimen. NMB D5845. Same specimen as Plate 21, figure 6. Calical surface, x5. . Figured Specimen. NMB D5842. Same specimen as Plate 21, figure 2. Calical surface, х 5. . Figured Specimen. NMB 105848. Upper Miocene, NMB locality 16911. Rio Mao, ?Gurabo Formation, Dominican Republic. Transverse thin-section, x25, showing one complete corallite. . Figured Specimen. NMB D5849. Upper Miocene, NMB locality 15846, Rio Gurabo, Gurabo Formation, Dominican Republic. Longitudinal thin-section, x 25, showing the wall between two corallites. BULLETIN 325 EXPLANATION OF PLATE 23 Poritesmacdonaldussauchan ees ea c I. uL Тт 78 SEM photographs of calices. Calices are fairly shallow with poorly developed pali and a prominent columella tubercle. The columellar synapticular ring is wide, and the septa do not always extend completely to it. Figure 1. Figured Specimen. NMB D5847. Same specimen as Plate 21, figure 8, x28. 2. Figured Specimen. NMB D5847. Same specimen as figure 1 above, x67. 3. Figured Specimen. NMB D5850. Lower Pliocene, NMB locality 15824, Río Gurabo, Mao Formation, Dominican Republic, x28. 4. Figured Specimen. NMB D5851. Upper Miocene, NMB locality 16883, Río Gurabo, Gurabo Formation, Dominican Republic, x 28. BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 23 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 24 b nr .. AN DOMINICAN REPUBLIC NEOGENE. 3: FOSTER EXPLANATION OF PLATE 24 Porites;portoricensis;GVaughan)4 Lac Rl 22. Branch fragments and thin-sections cut perpendicular to branch axes. The colony morphology is highly variable, ranging from thin branches to columns with skirts. Only one relatively complete colony has been found. Thin-sections cut perpendicular to each branch axis show a porous reticulum at the core of the axis, which is surrounded by uniform, prominent, widely-spaced horizontal septal junctions that may be crossed by vertical trabeculae. Figure IL. 2 3 Holotype. USNM 325061. Lower Miocene, USGS locality 3191, Lares Formation, Puerto Rico. Branch surface, x1. Figured Specimen. PIU WI9. Lower Miocene, Anguilla Formation, Anguilla. Branch surface, x1 (paratype of Goniopora clevei Vaughan). Figured Specimen. USNM 325115. Lower Miocene, USGS locality 6966, Anguilla Formation, Anguilla. Branch surface, x1 (paratype of Goniopora clevei Vaughan). . Figured Specimen. USNM 325072. Middle Miocene, USGS locality 6020, La Boca Formation, Panama. Branch surface, x 1 (holotype of Goniopora cascadensis Vaughan). . Figured Specimen. USNM 68315. Lower Miocene, USGS locality 6546, Tampa Formation, Florida. Branch surface, x1 (holotype of Goniopora matsoni Weisbord). . Possible synonym, lectotype of Porites (Synaraea) howei Vaughan. USNM 72890. Middle Miocene, USGS locality 6016, La Boca Formation, Panama. Branch surface, x1. . Figured Specimen. NMB D5796. Upper Miocene, NMB locality 15851, Rio Gurabo, Gurabo Formation, Dominican Republic. Branch surface, х1. . Figured Specimen. BM(NH) R28828. Neogene, Nivajé Shale, Dominican Republic. Branch surface, х1 (Duncan’s specimen of Alveopora fenestra Dana). . Figured Specimen. NMB D5797. Upper Miocene, NMB locality 15850, Rio Gurabo, Gurabo Formation, Dominican Republic. Branch surface, х1. . Figured Specimen. NMB D5798. Lower Pliocene, NMB locality 15821, Río Gurabo, Mao Formation, Dominican Republic. Branch surface, x 1. . Figured Specimen. ММВ 105799. Upper Miocene, NMB locality 16883, Rio Gurabo, Gurabo Formation, Dominican Republic. Branch surface, х1. . Figured Specimen. NMB D5800. Lower Pliocene, NMB locality 16884, Río Cana, Mao Formation, Dominican Republic. Branch surface, x1. . Figured Specimen. NMB D5802. Upper Miocene, NMB locality 15853, Río Gurabo, Gurabo Formation, Dominican Republic. Thin-section, x5. . Figured Specimen. NMB D5803. Upper Miocene, NMB locality 15855, Río Gurabo, Gurabo Formation, Dominican Republic. Thin-section, x5. . Figured Specimen. ММВ D5801. Upper Miocene, NMB locality 16933, Rio Gurabo, Gurabo Formation, Dominican Republic. Thin-section, х5. 103 BULLETIN 325 EXPLANATION OF PLATE 25 Porites portorscensis (Vaughan) m. wer... 2222-2504 79 Close-ups of calical surfaces. Calices vary widely in many characters; however, they are generally large and separated by an extensive coenosteal reticulum. Septal denticles are frequently well-developed. Figure 1. 2 3 4. Holotype. USNM 325061. Same specimen as Plate 24, figure 1. Calical surface, x5. Figured Specimen. USNM 25072. Same specimen as Plate 24, figure 4. Calical surface, x5. Figured Specimen. PIU WI10. Lower Miocene, Anguilla Formation, Anguilla. Calical surface, x 5 (holotype of Goniopora clevei Vaughan). Figured Specimen. PIU WI9. Same specimen as Plate 24, figure 2. Calical surface, х5. . Figured Specimen. USNM 68315. Same specimen as Plate 24, figure 5. Calical surface, x5. . Figured Specimen. USNM 325113. Same specimen as Plate 24, figure 6. Calical surface, x5. PLATE 25 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 ARA Mayr ҮЗ Ф; Ж," 4 7 : e ` t» ем d 222 4 5 LJ h а x on iN] > = 599%, 2 Ev Уча s, 8 ` ve BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 26 DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 105 EXPLANATION OF PLATE 26 Porites portoricensis NAME е ce eee o um 22 d. a al 79 Close-ups of calical surfaces and thin-sections. As in Plate 25, calices appear variable in many characteristics but are generally large and separated by a well-developed coenosteal reticulum. Thin-sections show the irregular development of vertical trabeculae, the prominent denticles, and the most common wall structure, which consists of three mural trabecular rows. Figure IL. 2 3. Figured Specimen. NMB D5797. Same specimen as Plate 24, figure 9. Calical surface, х 5. Figured Specimen. NMB D5800. Same specimen as Plate 24, figure 12. Calical surface, x5. Figured Specimen. NMB D5796. Same specimen as Plate 24, figure 7. Calical surface, х5. . Figured Specimen. NMB D5798. Same specimen as Plate 24, figure 10. Calical surface, x5. . Figured Specimen. NMB D5801. Same specimen as Plate 24, figure 15. Transverse thin-section, х 25, showing one corallite. . Figured Specimen. NMB D5804. Upper Miocene, NMB locality 16883, Río Gurabo, Gurabo Formation, Dominican Republic. Longitudinal thin-section, x 25, showing mural trabeculae between two corallites. Porites portoricensis (Vaughan) BULLETIN 325 EXPLANATION OF PLATE 27 SEM photographs of calices. Septa commonly bifurcate to give a Goniopora-like appearance. They are fre- quently formed by horizontal flakes with prominent denticles. Pali are weakly-developed; however, a small columella tubercle is usually present. Figure 1. 21 3 Figured Specimen. NMB D5805. Upper Miocene, NMB locality 16811, Río Gurabo, Gurabo Formation, Dominican Republic, х 28. Figured Specimen. NMB D5805. Same specimen as in figure 1 above, x67. Figured Specimen. NMB D5806. Upper Miocene, NMB locality 16934, Río Gurabo, Gurabo Formation, Dominican Republic, x 28. . Figured Specimen. NMB D5807. Upper Miocene, NMB locality 16883, Río Gurabo, Gurabo Formation, Dominican Republic, x 28. BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 27 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 28 DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 107 EXPLANATION OF PLATE 28 Porites portoricensis (V ogham) а ЕИ 2 79 SEM photographs of calices. In more worn material, the flaky texture of the septa is less apparent and the coenosteal reticulum dominates the surface morphology. Figure 1. Figured Specimen. NMB D5808. Upper Miocene, NMB locality 15851, Río Gurabo, Gurabo Formation, Dominican Republic, x28. 2. Figured Specimen. NMB D5808. Same specimen as figure 1 above, x67. 3. Figured Specimen. NMB D5809. Lower Pliocene, NMB locality 15821, Río Gurabo, Mao Formation, Dominican Republic, x28. 4. Figured Specimen. NMB D5810. Lower Pliocene, NMB locality 16884, Río Cana, Mao Formation, Dominican Republic, x28. 108 BULLETIN 325 EXPLANATION OF PLATE 29 Page Borueswaylandp noni nova на cun у есини MM LI a T M Nor M A ы E 81 Whole colonies. Colony morphology is massive, ranging in shape from columns to hemispherical mounds to plates. Figure 1. Holotype. USNM 325063. Middle Miocene, USGS locality 6016, La Boca Formation, Panama. Upper surface of a plate, x1. 2. Figured Specimen. USNM 66220. Lower Miocene, Ballast Point, Tampa Formation, Florida. Side of a nodular column, x1 (one of Wesibord's nontype specimens of “Porites floridaprima Bernard"). 3. Figured Specimen. NMB D5824. Upper Miocene, NMB locality 16811, Río Gurabo, Gurabo Formation, Dominican Republic. Upper surface of a plate, x». 4. Figured Specimen. NMB D5825. Upper Miocene, NMB locality 16933, Rio Gurabo, Gurabo Formation, Dominican Republic. Side of a smooth column, x1. BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 29 PLATE 30 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 109 EXPLANATION OF PLATE 30 Portteswaylandisnomanowe ge cer. Xe ш T Е о с ue 81 Close-ups of calical surfaces, thin-sections, and X-radiographs of colonies. This species is distinguished by a thick, porous wall, poorly-developed pali, a small columella tubercle, and septa that extend completely to the columella synapticular ring. X-radiographs and thin-sections show that growth was approximately 2 to 3 mm per year and that synapticulae, dissepiments, and trabeculae around the columella are especially well-devel- oped. Figure 1. Holotype. USNM 325063. Same specimen as Plate 29, figure 1. Calical surface, x10. 2. Figured Specimen. USNM 66220. Same specimen as Plate 29, figure 2. Calical surface, x10. 3. Figured Specimen. NMB D5825. Same specimen as Plate 29, figure 4. Calical surface, x 10. 4. Figured Specimen. NMB D5827. Lower Miocene, NMB locality 17284, Rio Yaque del Norte, Baitoa Formation, Dominican Republic. X-radiograph, х1. 5. Figured Specimen. NMB D5826. Upper Miocene, NMB locality 16856, Rio Cana, Cercado Formation, Dominican Republic. X-radiograph, x1. 6. Figured Specimen. NMB D5826. Same specimen as figure 5. Transverse thin-section, x 25, showing one complete corallite. 7. Figured Specimen. NMB D5826. Same specimen as figure 5. Longitudinal thin-section, x 25, showing one complete corallite. 110 BULLETIN 325 EXPLANATION OF PLATE 31 PO OOP OE HOL ли Са С cee UN EIN E ea уе у Жк 81 SEM photographs of calices. Septa extend completely to the columellar synapticular ring, which connects a well-developed network of trabeculae. The pali and columella tubercle are poorly-developed or absent. Figure 1. Figured Specimen. NMB D5828. Upper Miocene, NMB locality 15846, Rio Gurabo, Gurabo Formation, Dominican Republic, x 28. 2. Figured Specimen. NMB D5828. Same specimen as figure 1 above, x67. 3. Figured Specimen. NMB D5826. Same specimen as Plate 30, figures 4, 5, and 7, x28. 4. Figured Specimen. NMB D5829. Lower Miocene, NMB locality 16943, Río Yaque del Norte, Baitoa Formation, Dominican Republic, x 28. BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 31 PLATE 32 = ж m 2 a = O > К Q О = О E Z O [sa] E| < es E « = |52 El 2 < ш © n А E шщ =| El =) e DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 111 EXPLANATION OF PLATE 32 Goniopora hilli Vaughan a e 2 70202000 m T UE RU OR Ң 83 Whole colonies and X-radiographs of colonies. Colonies generally have smooth, hemispherical mound shapes that may be as large as 30 to 50 cm in diameter. Growth bands occur regularly at intervals of 3 to 4 mm. Figure 1. Holotype. USNM 325058. Middle Miocene, USGS locality 6016. La Boca Formation, Panama. Upper surface of colony fragment, х1. 2. Figured Specimen. NMB 05852. Lower Miocene, NMB locality 16937, Rio Yaque del Norte, Baitoa Formation, Dominican Republic. X-radiograph, х1. 3. Figured Specimen. USNM 68317. Lower Miocene, USGS locality 2084, Tampa Formation, Florida. Side view of colony, x1 (holotype of Goniopora tampaensis Weisbord). 4. Figured Specimen. NMB D5853. Upper Miocene, NMB locality 15861, Rio Gurabo, Gurabo Formation, Dominican Republic. Upper surface of colony, x1. 5. Figured Specimen. NMB D5854. Upper Miocene, NMB locality 16933, Rio Gurabo, Gurabo Formation, Dominican Republic. X-radiograph, х1. 6. Figured Specimen. NMB D5855. Lower Pliocene, NMB locality 16818, Rio Cana, Gurabo Formation, Dominican Republic. Side view of large colony, x Va. H2 BULLETIN 325 EXPLANATION OF PLATE 33 Gontoporm ME Мано папа рле GV, EEA E T TEE OT ROGET Эди соно чи 83 Close-ups of calical surfaces. Calices are highly-elevated, large, and separated by a relatively-reduced mural reticulum of variable thickness. The columella tangle and pali are poorly-developed; therefore, the fossa is characteristically V-shaped. Figure i 2. 3 CA Holotype. USNM 325058. Same specimen as Plate 32, figure 1. Calical surface, x 5. Figured Specimen. USNM 68317. Same specimen as Plate 32, figure 3. Calical surface, х 5. Figured Specimen. USNM 72891. Middle Miocene, USGS locality 6016, La Boca Formation, Panama. Calical surface, х 5 (lectotype of Goniopora canalis Vaughan). . Figured Specimen. USNM 325081. Lower Miocene, USGS locality 6775, White Springs, Florida. Calical surface, x 5 (paratype of Goniopora jacobiana Vaughan). . Figured Specimen. NMB D5853. Same specimen as Plate 32, figure 4. Calical surface, х 5. . Figured Specimen. NMB D5855. Same specimen as Plate 32, figure 6. Calical surface, x 5. PLATE 33 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 34 DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 113 EXPLANATION OF PLATE 34 Goniopora hilli Vanchah састы ты о. Wow a ee ee an eg C 83 SEM photographs of calices. The columella structure is reduced, and the septa extend almost completely to the calice center. The pali are weakly-developed. The septa are thick with prominent denticles. Figure 1. Figured Specimen. NMB D5853. Same specimen as Plate 32, figure 4 and Plate 33, figure 5, x28. 2. Figured Specimen. NMB D5853. Same specimen as figure 1 above, x67. 3. Figured Specimen. NMB D5854. Same specimen as Plate 32, figure 5, x 28. 4. Figured Specimen. NMB D5856. Lower Pliocene, NMB locality 16818, Rio Cana, Gurabo Formation, Dominican Republic, x28. 114 BULLETIN 325 EXPLANATION OF PLATE 35 Goniopora hilli Vaughan and Goniopora imperatoris Vaughan. Thin-sections. Corallites of G. hilli appear larger with thick septa and indistinct, irregular columellae; whereas in G. imperatoris, the septa are narrow with prominent pali, thick columella tangles, and a pronounced, well-developed coenosteal reticulum. Figure Page 1 Сено Fed S neemen NMB DS SOE a cin ЫЛЫЫ me esed ettet Ard T TROU ire OR O ee бык. КАЛД ve TE eet E 83 Same specimen as Plate 32, figure 5. Transverse thin-section, x5. 236 AUTE КОШОК SpecmuenawNVIBI58365 е еее 83 Same specimen as Plate 34, figure 4. Longitudinal thin-section, x 25, showing columella (center), dissepiments, and synapticulae. 3. G mmperatoris: Bigured SCORER КИБ DS S62 а тант аа neha ira Nass I IT 85 Upper Miocene, NMB locality 16853, Rio Cana, Cercado Formation, Dominican Republic. Transverse thin-section, х5. A. E DOOR Bioured. Specie NMB Юба О: 85 Upper Miocene, NMB locality 16856, Rio Cana, Cercado Formation, Dominican Republic. Longitudinal thin-section, x 25, showing mural trabeculae (right), columella (left center), dissepiments and synapticulae. 5. G POOD, ЕДЕП Specimen: BMNH R209678 222222222222. O ы ыб ке Ск 85 Upper Oligocene, chert and marl of Antigua. Transverse thin-section, x 25 (holotype of Alveopora daedalaea regularis Duncan). б. ТО PAD OOS Коше SPE OTE BMD RIOD Sc ет жил. oe eror d E O sc жет 86 Upper Oligocene, marl of Antigua. Transverse thin-section, х 25 (holotype of Alveopora daedalaea minor Duncan). TL GE UDOT, ENGIN Ce Speerin. BININIEDSNSZSGAD es леу. ежен а tube echec Аы ысыкы ООА Ee 86 Upper Oligocene, marl of Antigua. Polished surface, x 10 (holotype of Alveopora microscopica Duncan). PLATE 35 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 . - Vet BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 36 DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 115 EXPLANATION OF PLATE 36 Gonioporaimperatoris Vaughan ел e ceu c Ed 85 Whole colonies, close-ups of calical surfaces, and X-radiographs of colonies. Like G. hilli, colonies form large, smooth, well-rounded mounds with growth bands at intervals of 3 to 4 mm. However, corallites are smaller and the coenosteal reticulum more pronounced. Figure It 2: 3; Holotype. USNM 325049. Middle Miocene, USGS locality 6016, La Boca Formation, Panama. Upper colony surface, x1. Figured Specimen. NMB D5859. Upper Miocene, NMB locality 16853, Río Cana, Cercado Formation, Dominican Republic. Side view of colony, х. Holotype. USNM 325049. Same specimen as figure 1 above. Calical surface, x 5. . Figured Specimen. NMB D5859. Same specimen as figure 2 above. Calical surface, x5. . Figured Specimen. NMB 105858. Same specimen as Plate 35, figure 4. X-radiograph, x1. . Figured Specimen. NMB D5862. Same specimen as Plate 35, figure 3. X-radiograph, x1. . Possible synonym, holotype of Goniopora decaturensis silicensis Vaughan. USNM 325026. Lower Miocene, USGS locality 3381, Chattahoochee Formation, Georgia. Calical surface, x5. 116 BULLETIN 325 EXPLANATION OF PLATE 37 (GONTODOV GAIT CVALONISAN A О ey uc E a e a A a ата арала oe ek, 85 SEM photographs of calices. Corallites have numerous thin septa that extend to a well-developed columella tangle defined by several rows of synapticulae. Figure E 2 a Figured Specimen. NMB D5862. Same specimen as Plate 35, figure 3 and Plate 36, figure 6, x28. Figured Specimen. NMB D5862. Same specimen as figure 1 above, x67. Figured Specimen. NMB 105858. Same specimen as Plate 36, figure 5, x28. . Figured Specimen. NMB D5861. Lower Miocene, NMB locality 16944, Río Yaque del Norte, Baitoa Formation, Dominican Republic, x 28. BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 37 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 90 PLATE 38 DOMINICAN REPUBLIC NEOGENE. 3: FOSTER 117 EXPLANATION OF PLATE 38 Goniopora calhounensis Weisbord and Alveopora tampae Weisbord. Whole colonies and close-ups of calical surfaces. Calices of G. calhounensis are notably large with a thick coenosteal reticulum, well-developed pali and thick synapticulae. The Alveopora has characteristic thecal pores and irregular septal spines. Figure Page Гбзсашвоипепі OOY De ^ ESULGEISESq aem Ii ЕН 87 | Middle Miocene, Chipola Formation, Florida. Calical surface, x5. 2 GecalhounensissEiguredispecimensNMBID5SSOS UNT HAM ER... Ae зет eere у» ЧИА AN 87 Upper Miocene, NMB locality 16921, Rio Gurabo, Gurabo Formation, Dominican Republic. Side view of colony, x1. 37G#calhounensis#FigütediSpecimena NMBIDSSCSER мын cc ee ee eee E EM orae pcc ie шо 87 | Same specimen as figure 2 above. Calical surface, x 5. AF ADTAMPALATO POL Pen US NM тере Date ПАИ a hee ТИКЕ л екы NA мата 88 Lower Miocene, USGS locality 2084, Ballast Point, Tampa Formation, Florida. Side view of colony, х1. Jodie tampaesbopolypeeUSNMI66224e ET БЕ ITA Rena A libs уен» чен ав FS ЖЕ 88 Lower Miocene, USGS locality 2115, Hillsborough Bay, Tampa Formation, Florida. Side view of colony, х1. ON APTAmpaertDO DO per SNIMEO O22 dui анкета тантана з (tlc, suman mabe Dae тег ecce perii ds ny ss ke ee M 88 Same specimen as figure 5 above. Calical surface, x5. 1A. атраг капек Specimen МЕЗ ZOO err ate НЕК I LO A es ы ы RS tee DRE EA E 88 Upper Oligocene USGS locality 8713, El Limon, Tabera Group, Dominican Republic. Colony surface, x 5. 118 BULLETIN 325 INDEX Note: Page numbers are in light face, plate numbers are in bold face type; principal discussion pages are in italics. Agaricia dominicensis Vaughan, Шо 49 Almy and Carrion- Torres CIOS) sl 56 Alveopora Blainville, 1830 ........................ 48,50,52,59,71,74,88 chiapanecae Frost and Langenheim, 1974 ........................ 88 daedalaea var. minor Duncans 1863 а. 86 daedalaea var. regularis Duncan, 1863 ............................ 86 jenestrata (Eamarek). Dana, 1846 nun ne en 48,79 fenestrata (Dana) of Duncan, 1863........................... 74,79,81 TMICHKOSCODIEA Duncan, SOI eases dee 86 tampae Weisbord, 1973... 389. 52,53,74,88 ?Alveopora daedalea var. regularis Duncan, 1863 ................. 85 AMNH (American Museum of Natural History, New York, NY, MOBS = ect NM MEE e M 49,76,81,82,85,87 ES rn em een DET 60,74,79,81,85,87 CROCUS Вау les an EORR Ode Ce PEE TE UAE E 60 Anguila Бошан A eer c uc E terse 79,81,85,87 anguillensis, ОРИЗ E ЕРЕ ее ОА. 60,74,79,82 улы осше А уы АРЕ жыр о ЕАУ 78 ANSP (Academy of Natural Sciences, Philadelphia, PA, U.S.A.) .. ue pct O 48,49,87 T cu Ms еда ааа M M MU MC M m 74,81,84,86,87 AMER OLIMAR OMe echoes 81,87 АТО жет s KM аел Баи EEEE 62 AUTO МЕ WEB, TIIT aera eee Eee Pret ra» 49 Aspidosiphon corallicola Sluiter, 1889... ee 77 astreoides, Porites ............. Т 56,58,62,73,78,79,80,82, 2 ÜSEREDIACS-DIRZIHENSIS, POLIS у... голы tee nee egin dae ATAR ECO NODO Dias tote editors Рн өзе ЕРІ» ore PREIS UIA йк ores MM E TU t су ыт 59,83,88 Austria, Niederleismear Enzefeld va... een 62 Babel e ee er 49 Boitoa BOREAM nennen en 52,81,83,85-87 DISTRO GOMIOUONA О а ee 60,74,79,81 baracoaensis, Porites ................ 16,17,18 ...... 50,51,52,53,54, 55,60,64,66,67,68,69,70,71,73,74,75,76,81,89 baracoaensismalanzasensis, POTES coreana end ia Jas sion 60 Baracoa Form ton een Seve san AGE ау кыйрат CRT 76 лара САРЫНЫНА Т ЕЕ кы кс E 15 baraconensis уат WIQLANZACNSIS; PORES maana oa mo de ee ess eri reri 15 Bart саат A e Eee bue Ae 49 Beni ClO OO) О e crece ere SGC а ШК ене 59 incen Go OU Же a os du nM TU T m d 56 Bernard (1903) : . 56,59,60,74,75,82,83 Bernard (1905) E. 56,59,60,63,74,75,82 КОШЕ МШ ater Meth ER er 59,82 BELO Ін ID s ee orcs tone иии RO RO 49 б ah) конш озш к чы снн ele oer esti, Тел Же да ы э 49 TOI SUADISUCAL лев act. een oe 3 НІШ ОШ er ee nee 83,88 BMP program МИ en н RIOT EET v rro 63 BMIDP program NC CLG O een senden 63,68 BM(NH) == Museum [Natural History], London, England, U.K.) Ben 48,49,81,82 Bold HOGG) sn 2220210... 74 ВО CLOG OV nee een nee 74 BOB BU I nee ee: 74 Bonden TOMANDO еее teehee ижаара ROUES iy 76 BOSS OSL E O 68 Brake OVO E A ол ешек O TG. 56,59 Brakel ОЖ ая олуу een me лы кҮн weh 58 branten ENN е USA 560:02:73177583 Brazile menses rem re een VEO DU e Laces: 56,73,73 Abrolhos-Atchipelagor E ics ustedes ION ке к... 62 AbrolhosReeise nee лн ыле т essen. 62 Fernando dor NOVOME е нао 62 Parahyba do NORE И 62 Pernambuco, Canderas Reef 62 Farinha, Maria 62 Recife Military Air Base 62 Cas ӘЛЕ л и л a men E UT Cairns (1982) ai: calabricae, Porites calhounensis, Goniopora California, Sane Diego Com Barrett CANYON 2222. 60 Galoosaliatchee КОШТАШ он [| canalis, Goniopora carrizensis, Porites 60,74 cascadensis, Соторота ле. 60,74,79,80,81 Eércado Еошпалопи a ата ее ran 52,76,77,81,83,87 Chattahoochee Formations fai... en Ы tet emai’ 86,87 Cheste a bs x тарала marsh аан 49 Сеа СОПСОО mee «ceci tie ET ne йы Sedi 49 CHEST об о А 62,71,76,77,83,85, Hs hapa e ca a AL CODON Gera reser RD OC CHIDO ВОО И И. tee TR oe CPLO ЖИД АУА КККК suse у. АДЕП 59,74,76 Christen Ele КЕЕ» RR ETATE OU 49 Cisne and RADO re соко, 56 clavaria, Porites 62,71 clevei, Goniopora соотсав секта асет. 7 3l СОС ОТ LM RAE collegniana, Porites .. compressa, Porites .... convivatoris, Porites COOKS WV p M EM EE copoyensis, Goniopora corallicola, Aspidosiphon Corallium, Doris STEALS ot Seba, LOO ee TS Cornelius Bar ОТД БП Le Coryell and Ohlsen (1929) Cosmoporites Duchassaing and Michelotti, 1866 .................. ШИС